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ALBERT R. MANN
LIBRARY
AT
CORNELL UNIVERSITY
CORNELL UNIVERSITY LIBRARY
924 063 069 623
DATE DUE
OK.
600
M94 2-
Vv, #7
1993
00
MNIA*VN
Vi 4-7
m= MY COTAXON
AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS
Vol. XLVII April-June 1993
New and interesting Polypores from Mexico. ...... Leif Ryvarden and Gast6n Guzman 1
Glomales of Taiwan: III. A Compartative study of spore ontogeny in Sclerocystis
ear eae, COneee ol aes rk Gea doo he ee le ewe Chi-Guang Wu —s25
Systematic and biological studies in the Balansieae and related anamorphs.
III. Ascospore and macroconidial germination as a taxonomic criterion.
Ryan A. Phelps and G. Morgan-Jones 41
Chaetospermum chaetosporum (Coelomycetes). First record from the Iberian
Pees Soi i eee M. Muntanola-Cvetkovic and A. G6mez-Bolea 59
Lectotypification of Ophiobolus trichellus (Dothideales, Ascomycetes).
Christian Scheuer — 67
Taxomyces andreanae, a proposed new taxon for a bulbilliferous hyphomycete
associated with Pacific Yew (Taxus brevifolia).
Gary Strobel, Andrea Stierle, Don Stierle and W. W. Hess = 71
Two new South American species of Corticiaceae (Basidiomycetes) with amyloid
RRR eke) SS ee Wee Kurt Hjortstam and Leif Ryvarden = 81
Dematiaceous Hyphomycetes on Freycinetia (Pandanaceae). 4. Cryptophiale.
E. H. C. McKenzie and A. J. Kuthubutheen = 87
Genetics of a Tuber aestivum population (Ascomycotina, Tuberales).
G. Pacioni, G. Frizzi, M. Miranda and C. Visca 93
New species in the lichen family Parmeliaceae (Ascomycotina) from Australia.
John A. Elix 10]
Taxonomic studies in the genus Mycosphaerella. 2. Notes on some additional
Series OeCitisie On DIARsICHCCaG cS ee yy a we ke ks Michael Corlett 131
Development of conidiomata in the Phyllosticta state of Guignardia mangiferae Roy
and observations on the fine structure of the conidium.
J. Muthumary, J. A. Jayachandra and M. Bhagavathy Preetha 147
Hydnangium pila Pat., an older name for Martellia mediterranea Moreno, Galan
ee RR CCCN a ee a has haste G. Moreno and R. Galan 157
Computer coding of strain features of the genus Rhizopus.
Shung-Chang Jong and Candace McManus 161
Three new hyaloscyphaceous fungi from Norway and Greenland.
Sigurd Olsen, John Haines and Sigmund Sivertsen —_177
Taxonomic history of the oomycete genus Thraustotheca.
Will H. Blackwell and Martha J. Powell 183
[Contents continued overleaf]
ISSN 0093-4666 MYXNAE 47:1-508 (1993)
Published quarterly by MYCOTAXON, LTD., P. O. Box 264, Ithaca, NY 14851.
For subscription details, availability in microfilm and microfiche,
and availability of articles as tear sheets, see back cover.
[Contents continued from front cover]
Biostatistical characterization of the genus Oidium ................ Marco T. Ialongo
Hypogeous mycorrhizal fungi of Spain.
Isabel F. Alvarez, Javier Parladé, James M. Trappe and Michael A. Castellano
Three xylariaceous fungi with scolecosporous conidia.
Y..-M. Ju, Felipe San Martin Gonzalez and Jack D. Rogers
Biscogniauxia and Camillea in Mexico
Felipe San Martin Gonzalez and Jack D. Rogers
New fungi from Yucca: Planistromella yuccifoliorum, gen. et sp. nov., and its anamorph
Kellermania yuccifoliorum, sp. nov. and Planistromella uniseptata, sp. nov., the
teleomorph Of Kellermanid yuccipend. othe ace aie ota Annette W. Ramaley
4-O-Methyllividic acid, a new lichen depsidone. .. John A. Elix and Debra A. Venables
Anaeromyces, and earlier name for Ruminomyces.
Y. W. Ho, D. J. S. Barr, N. Abdullah, and S. Jalaludin
A new species of Piromyces from the rumen of deer in Malaysia.
Y. W. Ho, D. J. S. Barr, N. Abdullah, S. Jalaludin, and H. Kudo
Type specimens of lichens and lichenicolous fungi in the Canadian Museum
OF Nature (CANE) ea5 oie oo LE ere ek ae eee see Pak Yau Wong
Neosartorya primulina, a new species of food-borne ascomycetes.
Shun-ichi Udagawa, Noritsuma Toyazaki and Haruo Tsubouchi
Studies on Boleiss SeCUOWLURIAE! All boas yy Memes Sean eu ei eo akg R. Treu
Two new Crinipellinae (Tricholomataceae: Marasmiae) from South America.
Roy E. Halling
Septoria thymi sp. nov. from Bulgaria. . Ekaterina F. Sameva and Ganka G. Bakalova
Three new species and a new variety of Pluteus from the United States.
Partha Banerjee and Walter J. Sunberg
World list of cetrariowschens. 2.9 aol hn ware ee Tiina Randlane and Andres Saag
Two new species of Cristinia (Basidiomycotina, Aphyllophorales) and a survey
OF ThE SOnus 0). eee Kurt Hjortstam and Helga Grosse-Brauckmann
Comments on recent work on Ophiostoma and its synnematous anamorphs.
H. P. Upadhyay
Trematosphaeria pachycarpa and Herbampulla crassirostris gen. et spec. nov.
(Ascomycetes): Ti taclar eae ine cette naan aren amet te C. Scheuer and A. Nograsek
New or interesting lichenicolous fungi. 3. Karsteniomyces llimonae sp. nov. and
Sclerococcum serusiauxii sp. nov. (Deuteromycotina).
Montserrat Boqueras and Paul Diederich
Amanita reidii - a new species from South Africa.
Albert Eicker, J. V. Van Greuning and Derek A. Reid
Phytophthora hibernalis and P. syringdé...........0 0c eee H. H. Ho and S. C. Jong
A new Cortinarius from Southeastern (Fexas. 2s ce. eee Orson K. Miller
Book Reviews. ssc 8 sis th noid Bias Sie Gee cee eee Cee te eer rene L. M. Kohn
Notice: A new computer program for the identification of lichen
SUDS(ATICES ..0 Wea seks Esther Mietzsch, H. Thorsten Lumbsch and John E. Elix
Pthor TND EA oe asl ie eee & SS tae ini ec ane ome a dee ine ty eae rs er we
INDEX to fungous and lichen taxa.” “ees tn oc a Ce cn a ie es, yy
Errata, Publication Dates, MYCOTAXON Volumes 45 (corrected), 46...............
Reviewers
oe BD WPS. Oe. ER ee) lene! Tene he Leh eS [si ee fe) 10 ee eo le we, selma! ile te) rene A! (Gg he) wl Gt ce (@ Ghirennee-e (ee ie a 6) Se ie whe is.
MY COTAXON
AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS
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Published by
MYCOTAXON, LTD., P.O. BOX 264
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Printed in the United States of America
Table of Contents, Volume Forty-seven
April-June 1993
New and interesting Polypores from Mexico .. . Leif Ryvarden and Gaston Guzman
Glomales of Taiwan: III. A Compartative study of spore ontogeny in Sclerocystis
(eslOMaACCAGs, CSIOMALCS ) aa Nira tenis andere ae Pliers bee, Chi-Guang Wu
Systematic and biological studies in the Balansieae and related anamorphs.
II. Ascospore and macroconidial germination as a taxonomic criterion.
Ryan A. Phelps and G. Morgan-Jones
Chaetospermum chaetosporum (Coelomycetes). First record from the Iberian
BOUIN SU Ab we: Peua Pr iene, M. Muntanola-Cvetkovic and A. G6mez-Bolea
Lectotypification of Ophiobolus trichellus (Dothideales, Ascomycetes).
Christian Scheuer
Taxomyces andreanae, a proposed new taxon for a bulbilliferous hyphomycete
associated with Pacific Yew (Taxus brevifolia).
Gary Strobel, Andrea Stierle, Don Stierle and W. W. Hess
Two new South American species of Corticiaceae (Basidiomycetes) with amyloid
SDOLC Srunteeric Malet Alec haha deel ee oe Kurt Hjortstam and Leif Ryvarden
Dematiaceous Hyphomycetes on Freycinetia (Pandanaceae). 4. Cryptophiale.
E. H. C. McKenzie and A. J. Kuthubutheen
Genetics of a Tuber aestivum population (Ascomycotina, Tuberales).
G. Pacioni, G. Frizzi, M. Miranda and C. Visca
New species in the lichen family Parmeliaceae (Ascomycotina) from Australia.
John A. Elix
Taxonomic studies in the genus Mycosphaerella. 2. Notes on some additional species
DCCUUIAEFONL EST ASSICACCACI Gnas ahatner ail an ements Mee Ae A Michael Corlett
Development of conidiomata in the Phyllosticta state of Guignardia mangiferae Roy
and observations on the fine structure of the conidium.
J. Muthumary, J. A. Jayachandra and M. Bhagavathy Preetha
Hydnangium pila Pat., an older name for Martellia mediterranea Moreno, Galan
SSN LONIECEIN Gee Pe veeae Ch tee econ Meee gree te it G. Moreno and R. Galan
Computer coding of strain features of the genus Rhizopus.
Shung-Chang Jong and Candace McManus
Three new hyaloscyphaceous fungi from Norway and Greenland.
Sigurd Olsen, John Haines and Sigmund Sivertsen
Taxonomic history of the oomycete genus Thraustotheca.
Will H. Blackwell and Martha J. Powell
Biostatistical characterization of the genus Oidium.............. Marco T. Ialongo
my pogeous mycorrhizal fungi of Spain..................... Isabel F. Alvarez,
Javier Parladé, James M. Trappe and Michael A. Castellano
Three xylariaceous fungi with scolecosporous conidia.
Y..-M. Ju, Felipe San Martin Gonzalez and Jack D. Rogers
Biscogniauxia and Camillea in Mexico
Felipe San Martin Gonzalez and Jack D. Rogers
New fungi from Yucca: Planistromella yuccifoliorum, gen. et sp. nov., and its anamorph
Kellermania yuccifoliorum, sp. nov. and Planistromella uniseptata, sp. nov., the
teleomorph of Kellermania yuccigena................. Annette W. Ramaley
itt
25
4]
oy
67
701
101
iv
4-O-Methyllividic acid, a new lichen depsidone.
John A. Elix and Debra A. Venables
Anaeromyces, and earlier name for Ruminomyces.
Y. W. Ho, D. J. S. Barr, N. Abdullah, and S. Jalaludin
A new species of Piromyces from the rumen of deer in Malaysia.
Y. W. Ho, D. J. S. Barr, N. Abdullah, S. Jalaludin, and H. Kudo
Type specimens of lichens and lichenicolous fungi in the Canadian Museum
ob Nature (CANT) Sante cre mee ae Rien Mt Goel arises ane Pak Yau Wong
Neosartorya primulina, a new species of food-borne ascomycetes.
Shun-ichi Udagawa, Noritsuma Toyazaki and Haruo Tsubouchi
Studies’on Boletus \SectiOndUunidii anne eae aoe ke eis SoA i ed ete R. Treu
Two new Crinipellinae (Tricholomataceae: Marasmiae) from South America.
Roy E. Halling
Septoria thymi sp. nov. from Bulgaria.
Ekaterina F. Sameva and Ganka G. Bakalova
Three new species and a new variety of Pluteus from the United States.
Partha Banerjee and Walter J. Sunberg
World hst of céetranoid lichenss iano es eee Tiina Randlane and Andres Saag
Two new species of Cristinia (Basidiomycotina, Aphyllophorales) and a survey
Of the SenuUS aus oer We enc Ann Kurt Hjortstam and Helga Grosse-Brauckmann
Comments on recent work on Ophiostoma and its synnematous anamorphs.
H. P. Upadhyay
Trematosphaeria pachycarpa and Herbampulla crassirostris gen. et spec. nov.
C(ASCOMYCELES) 7 AMR nO da erie een Tak C. Scheuer and A. Nograsek
New or interesting lichenicolous fungi. 3. Karsteniomyces llimonae sp. nov. and
Sclerococcum serusiauxii sp. nov. (Deuteromycotina).
Montserrat Boqueras and Paul Diederich
Amanita reidii - a new species from South Africa.
Albert Eicker, J. V. Van Greuning and Derek A. Reid
Phytophthora hibernalis and P. syringae............. H. H. Ho and S. C. Jong
A new.Cortinarius from Southeastern Texas .....-.......0..0. Orson K. Miller
Book Revie wseanee tea ee hee trie eee Stee On at ot ft aeeaitane L. M. Kohn
Notice: A new computer program for the identification of lichen substances.
Esther Mietzsch, H. Thorsten Lumbsch and John E. Elix
Author, END Exar ids See dancin Sai oto eae nate is oe, ae geen fer TUG W ihre ean Sete ae
INDE X*torTangous and miche pn taxkarge eager men, er hee eee er oon eee ee
Errata, Publication Dates, MYCOTAXON Volumes 45 (corrected), 46...............
REVICWETS "GRA ue OP ee, Bararnd Gee ae BL ies Cee diel! SMR Oia re Rd Oem ana Me
MY COTAXON
Volume XLVI, pp. 1-23 April-June 1993
NEW AND INTERESTING POLYPORES FROM
MEXICO
by
Leif Ryvarden
UNIVERSITY OF OSLO
Box 1045, BLINDERN
N-0316 OSLO 3
NORWAY
and
Gaston Guzman
INSTITUTO DE ECOLOGIA
APARTADO POSTAL 63
XALAPA, VERACRUZ 91000
MEXICO
SUMMARY
Forty three species of polypores from Mexico studied by the authors
since 1989 are discussed, of which 19 are new records for the Mexican mycoflora
and 23 are reported from new localities. Most of the studied specimens are from
tropical vegetation, except a few from temperate regions or from coniferous
forests. The studied species are reported from the following states of the country:
Baja California, Distrito Federal, Jalisco, Morelos, Quintana Roo, Veracruz and
Yucatan.
RESUMEN
Se discuten 43 especies de Poliporaceos de México estudiadas por los
autores desde 1989, de las cuales 19 son nuevos registros para la micoflora
mexicana y 23 se presentan de nuevas localidades. La mayoria de los
especimenes proceden de vegetacién tropical, excepto unos pocos que son de
zonas templadas 0 de bosques de coniferas. Las especies consideradas se
adscriben a las entidades federativas: Baja California, Distrito Federal, Jalisco,
Morelos, Quintana Roo, Veracruz y Yucatan.
INTRODUCTION
The polypores of Mexico have been studied during the last
century, through the works of Kickx (1841) to Saccardo (1888), and
in the present centuary through several contributions by Murrill
(Murrill, 1905, 1907-1908, 1912, 1915), and modern revisions of
Anell and Guzman (1988), Ojeda-L6pez et a/. (1986), Valenzuela
and Nava (1991), Valenzuela and Chacdon-Jiménez (1991) and
Vazquez and Guzman-Davalos (1991). Nevertheless the knowledge
of the group is still scanty, as shown by the revision on some
collections in XAL Herbarium. Among these the authors found 19
new records for the country, and 23 from new localities not only
from the tropics where the mycological studies are few but also
from temperate regions, where the mycology has a good
development.
The present paper is based on the study of more than 70
specimens of polypores of XAL Herbarium (mostly duplicates from
ENCB), with duplicates in O. All the material was studied by the
authors between 1989-1991. The nomenclature following in this
paper, in according to Gilbertson and Ryvarden (1986-1987).
CONSIDERED SPECIES
A total of 43 species of polypores were studied, of which 19
are new records from Mexico, and 23 are reported from new
localities. Tables 1 and 2 show the studied species. Phylloporia
spathulata is considered in connection with a discussion of P.
veraecrucis from a new locality.
Albatrellus subrubescens (Murr.) Pouz.
This is an unusual North American species, only known from
Arizona and Florida in the U.S.A., and recently reported from
Mexico, from the States of Michoacan (Diaz-Barriga et al., 1988)
and Chiapas, Mexico, Nuevo Leon and Veracruz (Valenzuela &
Nava, 1991). The studied material is characterized by the
basidiocarps with yellowish, smooth to squamulose pileus, pore
surface greenish yellow with irregular reddish-orange tints, context
cream colored and unchanging, and by the absence of clamp
connections and amyloid spores.
3
Material studied. STATE OF VERACRUZ, Region E of Cofre
de Perote, S of Tembladeras, El Revolcadero, Pinus forests,
Villarreal 1424.
Amauroderma camerarium (Berk.) Furt.
It is a widespread fungus in the neotropics (Furtado, 1981),
and recognized by its pale yellowish context. Welden et a/. (1979)
and Anell & Guzman (1987, 1988), reported this fungus from the
State of Veracruz. |
Material studied. STATE OF QUINTANA ROO, road to the
sea from the road Puerto Morelos to Tulum, near the deviation to
Vallarta, tropical forest, Lopez 1848.
Amylosporus campbellii (Berk.) Ryv.
This is a new record from Mexico. It is a pantropical species;
in America known from southern United States, besides of Bermuda
(Gilbertson & Ryvarden, 1986-1987), and the Caribbean zone.
Murrill (1915) described this species as Tyromyces graminicola
Murr. Usually the species grows in clusters of semistipitate to
fanshaped basidiocarps on the ground in subtropical or tropical
vegetation. The pileus is whitish to pale brownish, grayish brown or
cinnamon brown, irregularly and finely tomentose. The poroid
surface is almost concolorous with the pileus. Microscopically the
small warted and strongly amyloid spores are characteristic and
point to relationships with the Hericiaceae where these features are
common.
Material studied. STATE OF MORELOS, road Cuernavaca to
Cuautla, Canon de Lobos, km 14, tropical forest, Guzman 6777.
Anomoporia albolutescens (Rom.) Pouzar
This is a temperate species, known from Europe and North
America through the coniferous forests (Gilbertson & Ryvarden,
1986-1987). The Mexican collection is the first record to Mexico. The
thin resupinate, loosely attached, rhizomorphic and pale yellowish,
yellow or orange yellowish basidiocarps are typical in this fungus.
Welden et al. (1979) reported A. myceliosa (Peck) Pouzar (as
Poria myceliosa Peck) from Acayucan, Ver. a species with white
basidiocarp.
Material studied. STATE OF VERACRUZ, Region E of Cofre
de Perote, S of Tembladeras, El Revolcadero, Pinus forest,
Montoya 318.
4
Antrodia gossypina (Speg.) Ryv.
A new record from Mexico. This species has a wide
distribution in temperate zones, mostly in coniferous forests
(Gilbertson & Ryvarden, 1986-1987). It is recognized by its
resupinate thick, soft to cottony, white to whitish or whitish rose
basidiocarp, with a rhizomorphic margin and by the ellipsoid
spores, 4.5-6 x 2-3 um.
Material studied. STATE OF VERACRUZ, Municipio de Totula,
El Mirador, subtropical forest with Quercus, Ventura 16294.
Antrodia malicola (Berk. & Curt.) Donk
This species was known from Mexico (as Trametes malicola
Berk. & Curt.) from a report by Sharp (1948), without specific
locality. It is widespread in warmer or temperate zones, common in
North America, but rare in Europe and Africa. The basidiocarps are
resupinate, effused-reflexed or sessile, forming nodulose to
imbricate pilei, whitish or pale woody color, with tomentose to
agglutinated glabrous surface .
Material studied. STATE OF VERACRUZ, Los Tuxtlas Region,
near El Jical, tropical forest, Guzman 19702.
Antrodia vaillantii (Fr.) Ryv.
Lowe (1966) reported this species as Poria vaillantii (Fr.)
Lowe from Mexico, but without specific locality. It is a widespread
and conspicuous crusty fungus, very close to A. gossypina (see
that). Many reports of this fungus are based on sterile specimens
having a well developed rhizomorphic margin, common also in that
species. A. vaillantii has spores (5-8 x 3-4 um) larger than A.
gossypina.
Material studied. STATE OF YUCATAN, between Xkantam
and Progreso, plantation of henequen (Agave fourcroydes Lem.),
Guzman 21365.
Antrodiella angulatopora Ryv.
This species was recently described from Venezuela
(Ryvarden, 1987), and this is the second report of the fungus. The
basidiocarp is sessile, broadly attached, imbricate and semicircular;
the upper surface is whitish to ochraceous with narrow orangish
yellow zones, and the pore surface is pale yellowish in dry
conditions.
Material studied. STATE OF VERACRUZ, road Xalapa to
5
Tlapacoyan, near Atzalan, La Calavera, subtropical forest with
Quercus and Cyathea, Ventura 12958.
Antrodiella versicutis (Berk. & Curtis) Gilbn. & Ryv.
This is a neotropical fungus, first reported from Mexico. The
allantoid spores separate this species from others of the genus. The
studied specimens have sessile basidiocarps or short or long
Stipitate, fanshaped pileus with the upper surface glabrous,
ochraceous to whitish yellow; pore surface almost concolorous with
the pileus, pores 6-8 per mm. Murrill (1907-1908) described this
species as Coriolus ochrotinctellus Murr. \t is very interesting to
observe the apparently abundance of A. versicutis in the tropical
vegetation of the eastern part of Peninsula of Yucatan, and the
absence of records from other tropical regions of Mexico, which
suggest that this is a Caribbean and South American species, as
those discussed by Guzman (1986).
Material studied. STATE OF QUINTANA ROO, road Felipe
Carrillo Puerto to Vigia Chico, 20 km from Felipe Carrillo Puerto,
tropical forest, Guzman 20777. Road Puerto Morelos to Tulum,
near the deviation to Vallarta, tropical forest, Guzman 23830; Lopez
1815; 1882. Road Coba to Nueva Xcan, near the deviation to
Sajaca-Chen, tropical forest, Lopez 1925. N of Reserva de la
Biésfera de Sialan Kalan, Archeological zone of Mugil, tropical
forest, Guzman 23853.
Aurificaria luteo-umbrina (Rom.) Reid
Murrill (1907-1908) described this species as Pyroporellus
subblinteus Murr. from Mexico (from the coasts of the State of
Sonora). A. luteo-umbrina is common in America, from the SE of
the U.S.A. to Brazil, and in Australia, but unknown from Africa
(Ryvarden & Johansen, 1980; Gilbertson & Ryvarden, 1986-1987).
The basidiocarps are sessile on logs, rusty to yellowish brown and
brittle, up to 80 mm wide; the context is bright yellowish brown,
instantaneously staining red with KOH; the pore surface turns
brownish red or cinnamon brown when touch in fresh conditions.
The lack of setae and the globose to subglobose spores, that stain
olivaceous brown in KOH define the species.
Material studied. STATE OF QUINTANA ROO, N of Reserva
de la Bidsfera de Sialan Kalan, Archeological zone of Mugil, tropical
forest, Guzman 23862.
6
Coriolopsis floccosa (Jungh.) Ryv.
This species was reported by Ojeda-Lopez et al. (1986) from
the State of Guanajuato. It is a pantropical fungus, apparently more
common in the old tropics than in the neotropics. The basidiocarps
are sessile, broadly attached and semicircular, with brown to
grayish pileus, with black concentric zones in dry specimens. It may
be confused with C. rigida (see that); however this latter species
is more thinner and flexible, and with pale brown pore surface.
Material studied. STATE OF QUINTANA ROO, road Puerto
Morelos to Tulum, near the deviation to Vallarta, tropical forest,
Guzman 23950.
Coriolopsis rigida (Berk. & Mont.) Murr.
Murrill (1907-1908, 1912), Welden & Lemke (1961), Castillo
et al. (1969), Castillo & Guzman (1970), Guzman (1972) and Anell
& Guzman (1987) reported this species from the State of Nuevo
Leon, Colima and Veracruz. This is a very common fungus in the
neotropics, but it is widespread even in the Eastern U.S.A. The
basidiocarp is variable in habit, from effused or connate to imbricate
clusters (See discussion of C. floccosa).
Material studied. STATE OF YUCATAN, 5 km S of Xochel,
road to Huhi, near Sacabah, tropical forest, Guzman 23746.
Daedalea microsticta Cooke Figs. 4-5
This is a widespread neotropical fungus. It was known from
the States of Colima and Veracruz (Murrill, 1912; Guzman, 1972,
Anell & Guzman, 1987). Here it is recorded from the States of
Nuevo Leon and Quintana Roo. The basidiocarp is applanate, more
or less thick, sessile, light color; the pileus is ivory color in dry
conditions, and reach more than 120 mm wide. Some studied
specimens were found in ENCB as Daedalea elegans Spreng. :
Fr., a neotropical fungus widespread in Mexico, that differs from D.
microsticta for its white funnel shapped basidiocarp with a thin
pileus.
Material studied. STATE OF NUEVO LEON, Municipio
Santiago, La Chancaca, Castillo 211; El Cercado, Sepulveda 19:
Ramirez s.n., all in subtropical forests. STATE OF QUINTANA ROO,
road Vigia Chico to Felipe Carrillo Puerto, Rancho Yaras, tropical
forest, Guzman 23934.
vf
Datronia caperata (Berk.) Ryv.
A pantropical species reported from Mexico as Polyporus
caperatus Berk., Coriolopsis caperata (Berk.) Murr., and C.
caperatiformis Murr., from the States of Campeche, Colima,
Hidalgo, Quintana Roo, Tabasco and Veracruz by Murrill (1912,
1915), Guzman (1972, 1975), Welden et a/. (1979), and Anell &
Guzman (1987). The sessile, thin and flexible basidiocarp is very
variable in form, from applanate, dimidiate, conchate to flabelliform,
up to 50 mm wide. In wet and shady places the pileus becomes
almost smooth with a tomentum restricted only to certain zones, but
in drier and sunny localities, it is covered with a characteristic dark
tomentum in distinct zones and below the tomentum there are well
defined black zones. The pore surface is white, but in old and dried
specimens is pale brownish, grayish brown to deep brown; in fresh
conditions turns brown when touched.
Material studied. STATE OF MORELOS, highway Mexico to
Cuernavaca, near the deviation to Cuautla, subtropical forest with
Quercus, Guzman 18114. STATE OF QUINTANA ROO, N of
Reserva de la Bidsfera de Sialan Kalan, Archeological zone of
Mugil, tropical forest, Guzman 23869. |
Diplomitoporus lenis (Karst.) Gilobn. & Ryv.
A new report from Mexico, this is a common circumglobal
species in subtropical and temperate areas (Gilbertson & Ryvarden,
1986-1987), but rare in the tropics. Ryvarden and Johansen (1980)
reported only one specimen from Africa [as Antrodia lenis (Karst)
Ryv.]. The spores are small, allantoid to Iunate and the basidiocarp
is resupinate, white to whitish, forming large crusts under the logs.
Material studied. STATE OF QUINTANA ROO, N of Reserva
de la Bidsfera de Sialan Kalan, Archeological zone of Mugil, tropical
forest, Guzman 23871.
Flavodon flavus (KI.) Ryv.
This species has hitherto not been reported from America.
It is very common in Africa, Asia and Australia (Ryvarden &
Johansen, 1980) and otherwise shows the relationships between
tropical regions of Mexico and Africa, observed by Guzman et al.
(1989). The intense yellow to orangish yellow basidiocarps are very
conspicuous. The hymenophore is hydnoid to iripiciform and the
pileus turns red with KOH.
Material studied. STATE OF QUINTANA ROO, road Coba to
Nueva Xcan, near the deviation to Sajaca-chen, Lopez, 1941; 1947.
Fomitopsis melliae (Underw.) Gilbn. & Ryv. pals bates
It is a new report from Mexico. This is a North American
species (Gilbertson & Ryvarden, 1986-1987), and the collections
from Mexico represent the southern limit so far known today. The
basidiocarp is up to 80 mm wide, thin to thick, sessile with a pileus
surface ivory to tan or cinereous color, frequently glabrous and
azonate (the collection from Quintana Roo is faintly zonate and
thin); the pore surface is whitish to ochraceous and the context is
whitish to pale buff.
Material studied. STATE OF QUINTANA ROO, N of Reserva
de la Biésfera de Sialan Kalan, Chuyaxchen, Archeological zone of
Mugil, tropical forest, Guzman 24147. STATE OF VERACRUZ, N of
Xalapa, Rancho Lucas Martin, subtropical disturbed vegetation,
Guzman 1915 (this specimen was reported by Guzman, 1963-A, as
Daedalea ambigua Berk.).
Ganoderma coffeatum (Berk.) Furt. Figs. 6-7
This species also placed in Amauroderma by Murrill or
Humphreya by Steyaert (1972), was recently reported from Mexico
by Cappello & Hernandez (1990), Vazquez & Guzman-Davalos
(1991) and Valenzuela & Chac6on-Jiménez (1991) from the States of
Jalisco, Tabasco and Tamaulipas. It is a neotropical and
widespread fungus, very common in South America. Its light
colored context, the brown ferrugineous velutinous pileus and stipe,
the whitish pore surface and the coarsely ornamented spores with
a prominent reticulated pattern of coherent ridges, are distinct good
features of this species. Macroscopically this fungus resembles an
Amauroderma. Here it is reported for first time from the States of
Quintana Roo and Veracruz.
Material Studied. STATE OF QUINTANA ROO, N of Reserva
de la Biosfera de Sialan Kalan, Archeological zone of Mugil, tropical
forest, Guzman 23902. STATE OF VERACRUZ, Uxpanapa region,
tropical rain forest, Guzman 30467.
Grammothele fuligo (B. & Br.) Ryv.
This is a tropical corticoid fungus, reported as Porogramme
fuligo (B. & Br.) Pat. by Lowe (1963) from Mexico without specifing
locality. The species occurs exclusively on dead leaves or stems of
9
monocotyledoneous, specially on all types of palms. It is easy to
overloook due to its resupinate basidiocarp with dark gray bluish
paints.
Material studied. STATE OF QUINTANA ROO, road to Xcan,
15 km from Coba, tropical forest, Lopez 1913.
Incrustoporia nivea (Jungh.) Ryv.
This is a circumglobal temperate species. It was reported
from Mexico by Sharp (1948) as Polyporus semipileatus Peck,
without specifing locality. Murrill (1912) reported it from the State of
Morelos as Tyromyces semipileatus (Peck) Murr. It is known in
North America as Skeletocutis nivea (Jungh.) Keller (Gilbertson &
Ryvarden, 1986-1987). It presents white to cream color annual
basidiocarp, effused-reflexed to resupinate.
Material studied. STATE OF BAJA CALIFORNIA, road
Ensenada to Ojos Negros, NE of Agua Viva, Quercus forest,
Guzman 24302-A.
Inonotus hispidus (Bull. : Fr.) Karst.
This is a common fungus in the United States (Gilbertson &
Ryvarden, 1986-1987), characterized by its strongly hispid upper
surface and the pigmented spores. The species was known from
Mexico from the States of Baja California, Hidalgo, Mexico,
Michoacan, Nuevo Leon and Veracruz (Bandala et a/., 1992). In the
present paper it is reported from Jalisco.
Material studied. STATE OF JALISCO, 11 km NE of
Tamazula, Rio Contla, tropical disturbed vegetation, Guzman 8593.
Inonotus munzii (Lloyd) Gilbn.
A subtropical to xerophitic warm temperate American fungus,
easy to recognize from the other species of the genus by the
branched setigerous elements of the pileus and lack of setae. The
basidiocarps are sessile, frequently ungulate and bright yellowish
brown to reddish brown or cinnamon brown. The species was first
reported from Mexico by Guzman (1963-A, -B) in subdesert lands
from the north to the central regions of the country. Castillo &
Guzman (1970) reported it from the State of Nuevo Leon. It is
reported here for first time from Distrito Federal.
Material studied. DISTRITO FEDERAL, Mexico City, Santo
Tomas Colonia, Campus of the National School of Biological
Sciences, on a living Schinus molle L., Fanti 74.
10
Junghuhnia nitida (Pers. : Fr.) Ryv.
This species was reported from the State of Oaxaca by
Welden et al. (1979). It is a cosmopolitan fungus, recorded from
Northern Norway to Australia and New Zealand. It grows on any
type of dead wood of angiosperms. Curiously in Mexico it is only
known from tropical regions. The pinkish-cinnamon color of the
pore surface and the so common thick-walled incrusted cystidia are
characteristic features of this species (Gilbertson & Ryvarden, 1986-
1987).
Material studied. STATE OF QUINTANA ROO, near the road
Puerto Morelos to Felipe Carrillo Puerto, Archeological zone of
Chunyaxche, tropical forest, Guzman 20646. STATE OF
VERACRUZ, Los Tuxtlas Region, 3 km S of Montepio, tropical rain
forest, Guzman 7257.
Microporellus obovatus (Jungh.) Ryv.
This is a pantropical species reported from Mexico by
Cappello & Hernandez (1990), Vazquez & Guzman-Davalos (1991)
and Valenzuela & Chacon-Jiménez (1991) from the States of
Jalisco, Tabasco and Tamaulipas. The white to yellowish orange or
straw color, tomentose to velvety striate, round to reniform or
spathulate pileus, sometimes radially striate, laterally stipitate, with
a white to concolorous pore surface and with very small pores (6-8
per mm) are features that distinguish this peculiar species
(Gilbertson & Ryvarden, 1986-1987).
Material studied. STATE OF QUINTANA ROO, road Puerto
Morelos to Vallarta, San Matilde, tropical forest, Guzman 20539.
State of Veracruz, Municipio de Totutla, El Mirador, subtropical
forest, Ventura 17324. Los Tuxtlas Region, 7 km S of Montepio,
Biological Station of the University of Mexico, tropical rain forest,
Guzman 7160. 2 km SW of Xalapa, near Rio Coapexpan,
subtropical forest with Quercus and Liquidambar, Bandala 227.
Oligoporus fragilis (Fr.) Gilobn. & Ryv.
A new report for the State of Morelos. This species was only
known from the States of Guanajuato and Michoacan (Cifuentes et
al., 1990; Ojeda-Lopez et a/., 1986) [in both cases as Tyromyces
fragilis (Fr.) Donk]. It is a boreal fungus following pine trees in high
altitudes. It is easy to recognize by the white to buff basidiocarps
that turn reddish brown when bruised or dried.
11
Material studied. STATE OF MORELOS, Lagunas de
Zempoala National Park, near the principal lagoon, Pinus forest,
Guzman 3174.
Perenniporia ohiensis (Berk.) Ryv.
Welden & Lemke (1961) and Castillo et a/., (1969) reported
this fungus as Fomes ohiensis (Berk.) Murr. from the State of
Nuevo Leon. The large (up to 13 um long) thick-walled, dextrinoid
spores are the principal features of this American species, as well
as its small effused-reflexed or ungulate basidiocarps, with pileus
whitish or ivory to brown or blackish, glabrous and azonate or fairy
zonate, and almost concolorous with the pore surface and context
(Gilbertson & Ryvarden, 1986-1987).
Material studied. STATE OF QUINTANA ROO, road Puerto
Morelos to Tulum, near the deviation to Vallarta, Botanical Garden
of CIQRO, tropical vegetation, Guzman 23788. N of Reserva de la
Biosfera de Sialan Kalan, Archeological zone of Mogil, tropical
forest, Guzman 23872. STATE OF YUCATAN, 5 km S of Xocchel
road to Hohi, near Sacabah, tropical forest, Guzman 23753. 5 km
S of Conkal, road to Telchac, tropical forest, Guzman 23595.
Perenniporia tephropora (Mont.) Ryv.
A new record from Mexico. It is a pantropical species, known
in the State of Louisiana (U.S.A.), the Caribbean region and Africa
(Gilbertson & Ryvarden, 1986-1987; Ryvarden & Johansen, 1980),
and considered as Loweporus by Ryvarden or Fomitopsis
cubensis by Murrill. It is characterized by its basidiocarps, which
are resupinate, large and thick, crustose and perennial, grayish to
dark ochraceous or milk coffee color to ferrugineous brown. The
basidiocarps frequently grow on vertical surfaces of the trunks.
Material studied. STATE OF YUCATAN, road Merida to
Valladolid, 3 km from Tixpeual, tropical forest, Guzman 23636.
Phellinus chryseus (Lév.) Ryv.
A new record from Mexico. It is a rare neotropical species,
Originally described from Colombia (Ryvarden, 1981; Ryvarden &
Johansen, 1980). The fairly large setae and small hyaline
dropshaped basidiospores, define the species, besides its typical
brown ferrugineous context and pore surface, as well as the rimose
and zonate grayish black pileus. The Mexican collection represents
the northern limit so far known to this fungus. The studied specimen
12
was growing in a disturbed tropical vegetation.
Material studied, STATE OF JALISCO, Tamazula region, near
Agua Caliente, tropical deciduous forest, Guzman 12486.
Phellinus contiguus (Fr.) Pat.
It is a widespread species in the tropics and warmer rare of
Europe (Gilbertson & Ryvarden, 1986-1987). The large hyaline and
ellipsoid spores and tramal setae characterize the species. The
basidiocarp is perennial and resupinate, forming a thick, rusty
brown crust below the logs. It is reported for first time from Mexico
from tropical forests of the Peninsula of Yucatan.
Material studied. STATE OF QUINTANA ROO, N of Reserva
de la Bidsfera de Sialan Kalan, Archeological zone of Mugil, tropical
forest, Guzman 23882.
Phellinus ferrugineo-velutinus (Henn.) Ryv.
This is a rare resupinate neotropical species, new recorded
from Mexico, and characterized by small hyaline spores and
scattered setae (Gilbertson & Ryvarden, 1986-1987). Probably
overlooked as other resupinate Phellinus species, which are often
neglected because of the considerable difficculties presented in
their identification. Ph. ferrugineo-velutinus forms large and thick,
ferrugineous brown crusts under the logs, with a conspicous
mustard yellow color margin.
Material studied. STATE OF QUINTANA ROO, road Puerto
Morelos to Tulum, near the deviation to Vallarta, road to the sea,
tropical forest, Lopez 1849.
Phellinus grenadensis (Murr.) Ryv.
New record from Mexico. It is a wide-spread species in the
Caribbean zone and Central America. It belongs to the. complex
group of pileate species without setae and rather uniformely
ellipsoid spores (Gilbertson & Ryvarden, 1986-1987). The studied
specimen has a small basidiocarp, no more than 20 mm wide,
subungulate, rusty brown and with a crustose pileus.
Material studied. STATE OF QUINTANA ROO, Municipio
Lazaro Cardenas, S of Chinquilla, tropical disturbed vegetation in
a grassland, Guzman 20657.
Phellinus rhytiphloeus (Mont.) Ryv.
This is a rare tropical species, previously known only from
13
Brazil (Ryvarden & Johansen, 1980). It has solitary, dimidiate, thick,
applanate and semicircular basidiocarps, with the typical brown
cinnamon color. When cut it presents a distinct black line below the
pileus. It is reported for first time from Mexico.
Material studied. STATE OF QUINTANA ROO, road Puerto
Morelos to Tulum, near Puerto Morelos, tropical forest, Lopez 1319.
Phellinus rimosus (Berk.) Pilat Fig. 1
This is a complex species with a widespread distribution in
the tropical and subtropical regions of the world. It grows in Africa
(Ryvarden & Johansen, 1980), but its presense in the U.S.A. is
doubtful (Gilbertson & Ryvarden, 1987). It was reported from
Mexico as Fomes, Cooke, from several temperate and subtropical
localities (Bandala et a/., 1992). It is reported here for first time from
Yucatan. The perennial, ungulate basidiocarp, with brownish to
blackish and tomentose to rimose pileus, yellowish-brown pore
surface and lack of setae are characteristic in this species.
Material studied. STATE OF YUCATAN, Municipio Progreso,
between Chelen and Chuburna, tropical vegetation, Guzman
21373-B.
Phylloporia chrysita (Berk.) Ryv.
A pantropical species, recorded from Mexico from the State
of Tamaulipas (Valenzuela & Chacon-Jiménez, 1991) and from the
Uxpana region in the State of Veracruz (Welden et a/., (1979). In the
presented paper it is reported from a new locality from the State of
Veracruz. The sessile, thick, rusty-brown basidiocarps, with the
pileus covered by a thick and velvety spongy compressed
tomentum, as well as the thin context separated by a dark line from
the tomentum, are good features that define the fungus and
separate it from Ph. frutica (Berk. & Curt.) Ryv., also known from
Mexico as parasitic on several tropical trees (Guzman, 1977).
Material studied. STATE OF VERACRUZ, Municipio Minatitlan,
Rio Coachapa, El Remolino, tropical forest, Pérez Ortiz 894.
Phylloporia spathulata (Hook.) Ryv.
It is a pantropical species, known as Polyporus by Cooke,
Coltricia by Murrill and Polystictus s. auct., rather common
although often overlooked due to its small size and dark brown
color basidiocarps, growing on the ground often among leaves and
debris inside the tropical forests. It seems to be a parasitic fungus,
14
at least specimens from Colombia were attached to living roots, and
others from Mexico present a long pseudorrhiza.
This fungus was reported from Mexico from the States of
Quintana Roo and Yucatan by Guzman (1983) and from Veracruz
by Guzman and Guzman-Davalos (1984). The species is
considered here, in spite of that the studied material is from the
same above regions of the country, because it is closely related to
Ph. veraecrucis (see below), which was considered as synonym,
and actually as an independent species; both species have golden-
yellow to cinnamon brown stipitated basidiocarps. Ph. spathulata
has pileus 10-60 mm broad and spores 3-4 x 2-3 um, while Ph.
veraecrucis presents pileus up to 100 mm wide and spores 4-4.5
X 3-3.5 um, as shown in the study of the type at K.
Material studied. STATE OF QUINTANA ROO, road
Escarcega to Chetumal, 10 km N from Tomas Garrido, Guzman
20910; 23884. Road Puerto Morelos to Tulum, near the deviation
to Vallarta, Guzman 21003; 21065; 23958; Lopez 1826; 1831.
STATE OF YUCATAN, road Merida to Valladolid, between Ticopo
and Tahnek, Guzman 23680. 5 km S of Xocchel, near Sacabah,
Guzman 23694; 23710 (all the collections from tropical forests).
Phylloporia veraecrucis (Berk. ex Sacc.) Ryv.
This is an interesting fungus described as Polyporus
veraecrucis Berk. from the State of Veracruz in the last century
(Berkeley, 1867), based in a collection on roots of a tropical tree in
1854. It was validated and redescribed by Saccardo (1888). Murrill
(1905) studied the type at K and considered the species as
Phaeolopsis veraecrucis (Berk.) Murr. Ryvarden & Johansen
(1980) considered Ph. veraecrucis as synonym of Ph. spathulata
(see that), but recently, Ryvarden (1991) recognized this species
and presented a modern description based on a study of the type.
In the present paper this fungus is reported for the first time from
the State of Quintana Roo.
Material studied. STATE OF QUINTANA ROO, Cozumel
region, Coba, tropical rain forest Guzman 20729.
Piptoporus soloniensis (Dub. : Fr.) Pil.
A new record from Mexico. This is a rare species distributed
in the SW of the United States (Gilbertson & Ryvarden, 1986-1987).
The Mexican collection seems to be the southern limit in the
continent. The basidiocarp is similar to Laetiporus sulphureus
15
(Bull. : Fr.) Murr. (Polyporus pseudosulphureus Long is a
synonym of P. soloniensis), but the discussed species has not the
bright orange and yellow sulphur coloration. The clamp connections
in the generative hyphae are a distinct feature.
Material studied. STATE OF VERACRUZ, road Xalapa to
Tlapacoyan, Municipio Atzalan, La Calavera, subtropical forest with
Quercus, Ventura 17315 (Anell & Guzman, 1988, reported this
specimen as Laetiporus sulphureus).
Polyporus varius Fr.
A circumglobal fungus specially common in temperate zones,
but scattered in the tropics (Ryvarden & Johansen, 1980, did not
report this species from Africa). The stipitate light coloured
basidiocarps with a black stipe make it rather distinct in the genus
except for P. elegans Bull. : Fr., that differs in the lack of the
striations of the pileus. The young specimens present light coloured
basal stipe as most of the studied specimens. This species has
been recorded from the States of Chiapas, Michoacan, Mexico
Tamaulipas and Veracruz (Bandala et a/., 1992). Here it is reported
for first time from Morelos, Quintana Roo and Yucatan.
Material studied. STATE OF MORELOS, IV Mushroom
Exposition of Morelos, Guzman 21911. STATE OF QUINTANA
ROO, N of Reserva de la Bidésfera de Sialan Kalan, Archeological
zone of Mugil, tropical forest, Guzman 23873. Road Puerto Morelos
to Tulum, near the deviation to Vallarta, tropical forest, Guzman
21015; 21077. STATE OF YUCATAN, 5 km S of Xocchel, road to
Huhi, near Sacabah, tropical forest Guzman 23687.
Rigidoporus vinctus (Berk.) Ryv. var. cinerea (Bres.) Setliff.
Anew record from Mexico. The variety vinctus was reported
from different localities in the State of Veracruz (Lowe, 1963,
Guzman, 1972, Welden et a/., 1979, Anell & Guzman, 1987) as
Poria vincta (Berk.) Curt. (except in the last reference, where it was
cited as Rigidoporus). The var. cinerea differs from the typic by
the grey to dark brown to almost black pore surface (pale
ochraceous buff to light pinkish ochraceous in the latter), and by
the perennial widely effused or effused-reflexed basidiocarp, with
pores 6-12 per mm.
Material studied. STATE OF QUINTANA ROO, 20 km from
Felipe Carrillo Puerto, road to Vigia Chico, tropical forest, Guzman
20773-B.
16
Trichaptum biforme (Fr.) Ryv. Fig. 3
A very common fungus in Mexico in temperate and tropical
regions, frequently reported as Polyporus pargamenus Fr.
(Guzman, 1977). It is closely related with Trichaptum abietinus
(Dicks. : Fr.) Ryv. from which differs mainly in the more purplish
bluing color of the pore surface and larger basidiocarps. T. biforme
is reported here for first time from the State of Quintana Roo.
Material studied. STATE OF QUINTANA ROO, road Puerto
Morelos to Tulum, near the deviation to Vallarta, road to the sea,
tropical forest, Lopez 1830. STATE OF VERACRUZ, Uxpanapa
region, tropical rain forest, Guzman 30466.
Tyromyces fumidiceps Atk.
This species was known only from Eastern U.S.A. and
Canada (Gilbertson & Ryvarden, 1987). The light greenish color of
the pore surface in herbarium specimens is characteristic. The
Mexican material has laterally stipitate basidiocarp, and whitish to
brownish yellow or brown chocolate color, azonate and tomentose
pileus. Its habitat is always close to stream and rivers, where the
basidiocarps are found on drift wood and similar debris.
Material studied. STATE OF MORELOS, old road Mexico City
to Cuernavaca, 3 km S of Tres Marias, Colonia Atlixtac, near a river,
Pinus- Quercus forest with subtropical elements, Guzman 8474.
Tyromyces galactinus (Berk.) Lowe
A new record from Mexico. This common North American
species (Gilbertson & Ryvarden, 1986-1987) is recognized by its
soft, white to whitish gray or pale ochraceous, strigose to
tomentose basidiocarps, with duplex context and fragant odour.
The studied specimen presents imbricate basidiocarps growing on
a living trunk of Platanus lindeniana Mart. & Gal.
Material studied. STATE OF VERACRUZ, old road Xalapa to
Coatepec, deviation to Rancho Guadalupe, way to Instituto de
Ecologia, subtropical (mesophitic) disturbed forest, Bandala-Munoz
Py
Tyromyces humeana (Murr.) Lowe
The species was only known from the type locality in Florida
(U.S.A) (Gilbertson & Ryvarden, 1986-1987). The Mexican collection
agrees well with Murrill’s fungus, except by the anise odour in fresh
basidiocarps which was not indicated in the studied specimens. The
17
basidiocarp is white but in dry condition turns brownish even
cinnamon brown in the center of the pileus.
Material studied. STATE OF MORELOS, road El Parque to
Tepoztlan, near Tepozteco mountain, subtropical forest, Guzman
5451.
Table 1. Polypores new from Mexico
Amylosporus campbellii (Berk.) Ryv.
Anomoporia albolutescens (Rom.) Pouzar
Antrodia gossypina (Speg.) Ryv.
Antrodiella angulatopora Ryv.
A. versicutis (B. & C.) Gilbn. & Ryv.
Diplomitoporus lenis (Karst.) Gilbn. & Ryv.
Flavodon flavus (KI|.) Ryv.
Fomitopsis meliae (Underw.) Gilbn. & Ryv.
Perenniporia tephropora (Mont.) Ryv.
Phellinus chryseus (Lév.) Ryv.
Ph. contiguus (Fr.) Pat.
Ph. ferrugineo-velutinus (Henn.) Ryv.
Ph. grenadensis (Murr.) Ryv.
Ph. rhytiphloeus (Mont.) Ryv.
Piptoporus soloniensis (Dub. : Fr.) Pil.
Rigidoporus vinctus var. cinerea (Bres.) Selif.
Tyromyces fumidiceps Atk.
T. galactinus (Berk.) Lowe
T. humeana (Murr.) Lowe
18
19
Figs. 1-3.- 1: Phellinus gilvus (Kruse, s.n., from Guerrero). 2:
Fomitopsis melliae (Guzman 1915, from Veracruz). 3:
Trichaptum biforme (Guzman 30466 from Veracruz).
Figs. 4-7.-4-5: Daedalea microsticta (Sepulveda 19, from Nuevo
Leon). 6-7: Ganoderma coffeatum (Guzman 30467, from
Veracruz).
20
Table 2. Polypores from new localities in Mexico
atrellus subrubescens (Murr.) Pouz.
Amauroderma camerarium (Berk.) Furt.
Antrodia malicola (B. & C.) Donk
A. vaillantii (Fr.) Ryv.
Aurificaria luteo-umbrina (Rom.) Reid
Coriolopsis floccosa (Jungh.) Ryv.
C. rigida (Berk. & Mont.) Murr.
Daedalea microsticta Cooke
Datronia caperata (Berk.) Ryv.
Ganoderma coffeatum (Berk.) Furt.
Grammothele fuligo (B. & Br.) Ryv.
Incrustoporia nivea (Jungh.) Ryv.
Inonotus hispidus (Bull. : Fr.) Karst.
!. munzii (Lloyd) Gilbn.
Junghuhnia nitida (Pers. : Fr.) Ryv.
Microporellus obovatus (Jungh.) Ryv.
Oligoporus fragilis (Fr.) Gilbn. & Ryv.
Perenniporia ohiensis (Berk.) Ryv.
Phellinus rimosus (Berk.) Pilat.
Phylloporia chrysita (Berk.) Ryv.
Ph. veraecrucis (Berk. ex Sacc.) Ryv.
Polyporus varius Fr.
Trichaptum biforme (Fr.) Ryv.
ACKNOWLEDGEMENTS
The authors acknowledge the authorities of their institutions
for the facilites to their researches. The authors are grateful to Victor
M. Bandala and Fidel Tapia of the Mycological Collection of XAL for
his help in the herbarium and to Leticia Montoya from Instituto de
Ecologia for revising this paper. Dr. Ake Strid, from Museum of
Natural History, Stockholm, Sweden, kindly revised critically the text.
LITERATURE CITED
Anell, J.C. and G. Guzman, 1987. Especies de Poliporaceos citadas
del Estado de Veracruz. Rev. Mex. Mic. 3: 137-148.
21
Anell, J.C. and G. Guzman, 1988. Nuevos registros de los hongos
del grupo de los Poliporaceos del Estado de Veracruz. Fev.
Mex. Mic. 4 : 25-42.
Bandala, V.M., G. Guzman and L. Montoya, 1992. Los hongos del
grupo de los Poliporaceos conocidos en México.
Universidad Autonoma de Nuevo Leon, Linares (en prensa).
Berkeley, M.J., 1867. On some new fungi from Mexico. Linn. Soc.
Jour. (Bot.) 9: 423-425.
Cappello, S. and H. Hernandez, 1990. Lista preliminar de los
hongos (macromicetos) y mixomycetes de Tabasco, México.
Universidad y Ciencia 7 (13) : 15-21.
Castillo, J. and G. Guzman, 1970. Estudios sobre los Poliporaceos
de Nuevo Leon, Il. Bol. Soc. Bot. Mex. 31 : 1-47. .
Castillo, J., G. Guzman and G. Sepulveda, 1969. Estudios sobre los
Poliporaceos de Nuevo Leon. |. Ciencia, Mex. 27 : 9-18.
Cifuentes, J., M. Villegas, L. Perez-Ramirez, M. Bulnes, V. Corona,
M. del R. Gonzalez, |. Jiménez, A. Pompa and G. Vargas,
1990. Observaciones sobre la distribuciOn, habitat e
importancia de los hongos de Los Azufres, Michoacan. Fev.
Mex. Mic. 6: 133-149.
Diaz-Barriga, H., F. Guevara-Fefer and R. Valenzuela, 1988.
ContribuciOn al conocimiento de los macromicetos del
Estado de Michoacan. Acta Botanica Mex. 2: 21-44.
Furtado, J.S., 1981. Taxonomy of Amauroderma (Basidiomycetes,
Polyporaceae). Mem. N.Y. Bot. Gard. 34 : 1-109.
Gilbertson, R.L. and L. Ryvarden, 1986-1987. North American
Polypores. Fungiflora, Oslo (vols. 1 & 2).
Guzman, G., 1963-A. Frecuencia y distribucion de algunos
Basidiomycetes lignicolas importantes en Mexico. An. Esc.
Nac. Cienc. Biols. 12: 23-41.
Guzman, G., 1963-B. Macromicetos de las zonas aridas de Mexico,
|. An. Esc. Nac. Cienc. Biols. 12: 43-60.
Guzman, G., 1972. Macromicetos mexicanos en el Herbario The
National Fungus Collections de E.U.A. Bol. Soc. Bot. Mex.
322731-55.
Guzman, G., 1975. Hongos mexicanos (macromicetos) en los
herbarios del extranjero Ill. Bol. Soc. Mex. Mic. 9: 85-102.
Guzman, G., 1977. /Identificacién de los hongos. Ed. Limusa,
Mexico City.
22
Guzman, G., 1983. Los hongos de la Peninsula de Yucatan, Il.
Nuevas exploraciones y adiciones micolégicas. Biotica 8:
71-100.
Guzman, G., 1986. Distribuci6n de los hongos en la region del
Caribe y zonas vecinas. Caldasia 15 : 103-120.
Guzman, G., V.M. Bandala, L. Montoya and Y. Saldarriaga, 1989.
Nuevas adiciones sobre las relaciones micofloristicas entre
Africa y el neotropico. El genero Rugosospora Heinem.
(Fungi, Agaricales). Brenesia 32: 107-112.
Guzman, G. and L. Guzman-Davalos, 1984. Nuevos registros de
hongos en el Estado de Veracruz. Bol. Soc. Mex. Mic. 19:
221-244,
Kickx, J., 1841. Notice sur quelques champignons du Mexique. Bull.
Acad. Sc. Brux. 8: 72-81.
Lowe, J.L., 1963. A synopsis of Poria and similar fungi from the
tropical regions of the world. Mycologia 55 : 453-486.
Lowe, J.L., 1966. Polyporaceae of North America. The genus Poria.
State Univ. College of Forestry, Syracuse University, Tech.
Publ. 90, Syracuse.
Murrill, W.A., 1905. The Polyporaceae of North America. XIl. A
synopsis of the white and bright-colored pileate species. Bull.
Torrey Bot. Club. 32: 469-493.
Murrill, W.A., 1907-1908. Polyporaceae. /n: North American Flora 9.
N.Y. Bot. Garden, New York.
Murrill, W.A., 1912. The Polyporaceae of Mexico. Bull. New York
Bot. Gard. 8 (28) : 137-153.
Murrill, W.A., 1915. Tropical Polypores. Lancaster, New York.
Ojeda-Lopez, S., M. Sandoval and R. Valenzuela, 1986. Los
Poliporaceos de México. |. Descripcién de algunas especies
del noreste de Guanajuato. Rev. Mex. Mic. 2: 367-436.
Ryvarden, L., 1981. Type studies in the Polyporaceae 13. Speccies
described by J.H. Léveille. Mycotaxon 13: 175-186.
Ryvarden, L., 1987. New and interesting polypores from tropical
| America. Mycotaxon 28: 525-541.
Ryvarden, L., 1991. Genera of Polypores. Nomenclature and
Taxonomy. Synopsis Fungorum 5, Fungiflora, Oslo.
Ryvarden, L. and |. Johansen, 1980. A preliminary polypore flora of
East Africa. Fungiflora, Oslo.
Saccardo, P.A., 1888. Sylloge Fungorum. Vol. 6 (Reprint 1944,
Edwards Brothers, Ann Arbor).
25
Sharp, A.J., 1948. Some fungi common to the highlands of
Mexico, Guatemala and Eastern United States. Mycologia
52 : 856-876.
Steyaert, R.L., 1972. Species of the Ganoderma and related
genera mainly of the Bogar and Leiden Herbaria. Persoonia
(és her ike):
Valenzuela, R. and R. Nava, 1991. E/ género Albatrellus
(Polyporaceae) en México. Memorias IV Congreso Nacional
de Micologia, Tlaxcala.
Valenzuela, R. and S. Chacén-Jiménez, 1991. Los Poliporaceos de
Mexico. Ill. Algunas especies de la Reserva de la Biosfera El
Cielo, Tamaulipas. Rev. Mex. Mic. 7 : 39-70.
Vazquez, L.S. and L. Guzman-Davalos, 1991. Nuevos registros de
Poliporaceos estipitados de Jalisco. Rev. Mex. Mic. 7:
113-120.
Welden, A.L., L. Davalos and G. Guzman, 1979. Segunda lista de
los hongos, liquenes y mixomicetos de las regiones de
Uxpanapa, Coatzacoalcos, Los Tuxtlas, Papaloapan y Xalapa
(México). Bol. Soc. Mex. Mic. 13: 151-161.
Welden, A.L. and P.A. Lemke, 1961. Notas sobre algunos hongos
mexicanos. Bol. Soc. Bot. Mex. 26: 1-24.
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MY COTAXON
Volume XLVI, pp. 25-39 April-June 1993
GLOMALES OF TAIWAN:
Ill. A COMPARATIVE STUDY OF SPORE ONTOGENY IN
SCLEROCYSTIS
(GLOMACEAE, GLOMALES)!
CHI-GUANG WU
Soil Microbiology Lab, Agricultural Chemistry Department,
Taiwan Agricultural Research Institute, Wu-feng, Taichung,
Taiwan, R.O.C.
KEY WORDS: Sclerocystis, sporophore, sporocarp formation, spore
germination, spore ontogeny.
ABSTRACT
During the course of investigation on the Glomales of
Taiwan, numerous sporocarps of Sc/erocystis were
isolated and identified as SS. coremioides, S. clavispora, S.
sinuosa, S. rubiformis, S. taiwanensis, and S.
liquidambaris. After detailed comparison among these
species by optical and electron microscopy, the similarity of
spore ontogeny and spore arrangement in sporocarps was
determined. Chlamydospores of S. coremioides with long,
slender subtending hyphae also produced sporophores the
same as those of other species of Sclerocystis and Glomus.
The chlamydospores of S. coremioides are not always
occluded by septa. The suggestion to define Sc/erocystis as
a monotypic genus is rejected. A modified generic concept
and a hypothesis to explain sporocarpic evolution of
Sclerocystis are proposed.
1. Contribution No. 1637 from Taiwan Agricultural Research Institute.
26
INTRODUCTION
Glomus and Sclerocystis were considered as closely related
groups and Sclerocystis appeared to be evolutionarily advanced in
sporocarp morphology from G/omus (Gerdemann and Trappe, 1974).
Recently, Almeida and Schenck (1990) proposed to transfer most of the
species of Sclerocystis to Glomus and retained only S. coremioides Berk.
and Br. in the genus, because there were at least four uniquely different
features in S. coremioides, i.e. (i) each spore forms individually on a
subtending hypha without branching to form a sporophore near the spore
base, (11) spores have a well-defined septum, (iii) spores are arranged in a
hemispherical layer, and (iv) sporocarps form distally or laterally from the
older sporocarps and appear fused in a column or branch. Almeida and
Schenck (1990) concluded that these characters were not found in other
species of Sclerocystis and Glomus. They applied Madelin's (1970) mode
of sympodial conidial formation to distinguish spore ontogeny of Glomus
from Sclerocystis, and postulated that S. rubiformis Gerdemann and
Trappe (1974), S. clavispora Trappe (1977), S. sinuosa Gerdemann and
Bakshi (1980), S. faiwanensis Wu and Chen (1987), and S.
liquidambaris Wu and Chen (1987) should be transferred to Glomus.
They have the same type of spore ontogeny as found in G. ambisporum
Smith and Schenck, G. heterosporum Smith and Schenck (Smith and
Schenck, 1985), and other species of Glomus. Glomus species have the
ability to generate a lateral sporophore in the subtending hypha near the
spore base.
During the survey of Glomales of Taiwan, abundant sporocarps
of Sclerocystis were collected from a variety of sites. These materials
provided all of the presently described species of Sclerocystis and were
very beneficial to the taxonomic study. In this paper, new evidence of
spore ontogeny and sporocarp formation, a modified generic concept of
Sclerocystis, and a hypothesis of sporocarpic evolution are presented.
MATERIAL EXAMINATIONS
Sporocarps were isolated by a wet-sieving and decanting
(Gerdemann and Nicolson, 1963) and were observed by optical and
scanning electron microscopy. In order to observe the central plexus of
sporocarps, fruiting bodies were ultrasonicated to break down peridial
tissues and chlamydospores.
af
The following specimens were examined:
Sclerocystis clavispora-T AIWAN: Chi-tou, from rhizosphere of
Chimonobambusa quadrangularis (Fenzi) Markino, Wu-850821; Chi-
tou, from the rhizosphere of Erechthites valerianifolia (Wolf) DC., Wu-
8508201; Chi-tou, from the rhizosphere of Gingo biloba L., Wu-
8508202; Chi-tou, from the rhizosphere of Colocasia formosanum
Hayata, Wu-8607151; Kenting, from the rhizosphere of Colocasia
formosanum, Wu-8605171; Pintong, from the rhizosphere of asparagus
(Asparagus officinalis L.), Wu-860813.
Sclerocystis coremioides-T AIWAN: Tainan, from rhizosphere of
Ageratum conyzoides L., Wu-850725; Tainan, from the rhizosphere of
orange (Citrus sinensis Osb.), Wu-8608141; Taipei, from the rhizosphere
of Bignonia sp., Wu-850810; Chi-tou, from the rhizosphere of Colocasia
formosanum, Wu-8607152; Pintong, from the rhizosphere of asparagus
(A. officinalis), Wu-860813; USA: Oregon, F55365S.
Sclerocystis liquidambaris-T AIWAN: Taipei, Holotype, from
rhizosphere of Liguidambar formosana Hance, Wu-860203; Isotype,
Wu-860627; Kaoshiun, Liou-Kuei, from the rhizosphere’ of
Cunninghamia lanceolata (Lamb.) Hook, Wu-870117.
Sclerocystis rubiformis-T AIWAN: Chi-tou, from the rhizosphere
of Erechthites valerianifolia, Wu-8508203; Chi-tou, from the
rhizosphere of Gingo biloba, Wu-8508204; Chi-tou, from the
rhizosphere of Colocasia formosanum, Wu-8607153; Kenting, from the
rhizosphere of Colocasia formosanum, Wu-8605 172.
“Sclerocystis sinuosa-TAIWAN: Tainan, Anpin; from the
rhizosphere of Miscanthus sp. in Casuarina equisetiffolia L . woods,
July 25, 1985, Wu-850725; Tainan, Anpin, from the rhizosphere of
ground covers in C. equisetifolia woods, September 18, 1985, Wu-
850918; Pintong, by Kau-ping River, from the rhizosphere of legume
(Glysine sp.), August 14, 1986, Wu-8608142; Pintong, by Kau-ping
River; from the rhizosphere of asparagus (A. officinalis), August 13,
1986, Wu-860813; Pintong, Tongang, from the rhizosphere of
unidentified herb grass, August 13, 1986, Wu-ME2; Tainan, Charlu, from
the rhizosphere of citrus (Citrus sp.), August 14, 1986, Wu-8608143;
Nantou, Chi-tou, from the rhizosphere of Sinocalamus sp., August 21,
1985, Wu-8508144; Taichung, Wu-feng, from the rhizosphre of maize
28
(Zea mays L.), November 14, 1991, Wu-111401; Orchid Island,
Taitung, from the rhizosphere of C. equisetifolia, July 7, 1986, WU-
860707; U.S.A., INVAM collection, Gainesville, Florida (SSNS 122).
Sclerocystis taiwanensis-T AIWAN:Chi-tou, Holotype, from the
rhizosphere of Crassocephalum rabens (Juss. ex Jacq.) S. Moore, Wu-
8508205; Chi-tou, from the rhizosphere of Colocasia formosanum, Wu-
8607154; Taipei, from the rhizosphere of unknown host, Wu-850810;
Tainan, from the rhizosphere of orange (C. sinensis), Wu-850724.
RESULTS
SPOROPHORE OF SCLEROCYSTIS: Sporophores branching from the
base of chlamydospore are very commonly found in the every species of
Sclerocystis. The sporophores are of various shapes such as
membraneous vesicles (Figs. 1, 3, 4 ), dichotomous branches (Figs. 2, 6),
or irregular forms. Sporophores are not uncommon in S. coremioides.
They may be globular structures or tubular hyphal branches (Figs. 4, 7).
The formation of sporophores is by branching from the chlamydospores
with or without septa. Sporophores of S. /iqguidambaris are further
differentiated into clubshaped structures which are parts of the peridium
(Eigse oy
SEPTA OF CHLAMYDOSPORES: Septa are very often formed at the
bases of chlamydospores in Sclerocystis (Figs. 3, 4). The frequency of
septa depends on the species and represents one of character in the
species description. The septum is very commonly, but not always, found
below the base of chlamydospores in S. coremioides (Figs. 4, 7). A
Figures. 1-6. Micrographs of Sclerocystis spp. Fig. 1. Chlamydospores of
S. sinuosa with a sporophore (arrow head). Fig. 2. Chlamydospores of S.
liquidambaris with a sporophore (arrow head). Note paraphysis-like
structures (P) enclosing the chlamydospore. Fig. 3. Chlamydospores of S.
taiwanensis with sporophores (arrow head). Note the septa formed at the
bases of spores. Fig. 4. A sporophore of S. coremioides formed below
the spore base (arrow head). Note the presence of a membraneous
septum (arrow) at the spore base. Fig. 5. Mature (upper) and vesicular
young spores (YS) of S. clavispora. Note the mature spore enclosed by a
exospore wall (arrows). Fig. 6. Hyphal branches formed from the spore
base of S. clavispora. Scale= 30 tm.
29
30
similar type of septum occurs in S. sinuosa and S. taiwanensis (Fig. 3).
Septa are also present in the subtending hyphae of S. rubiformis;
however, they are much thinner than those of S. coremioides.
ARRANGEMENT OF CHLAMYDOSPORES IN SPOROCARPS: The
chlamydospores of S. coremioides are arranged in a hemispherical layer
(Figs. 10, 16). The intersporal spaces are traversed by hyphae which
grow radially from the central plexus and terminally connect to the
peridium. Similar intersporal hyphae occur in S. sinuosa and _ “S.
liquidambaris (Figs. 2, 9). In young sporocarps of S. rubiformis,
chlamydospores are also arranged in a hemispherical layer (Fig. 12).
However, with the subsequent formation of additional chlamydospores,
these immature sporocarps are transformed into globular sporocarps with
radially arranged chlamydospores.
CHLAMYDOSPORE AND SPOROCARP FORMATION: All of the
chlamydospores in sporocarps of Sclerocystis are produced from a
central plexus. The central plexus ranges from a single flat, stellate cell
as found in S. rubiformis and S. taiwanensis (Figs. 12, 15), to a mass of
interwoven hyphae as found in S. clavispora, S. liquidambaris, S.
sinuosa, and S. coremioides (Figs. 8, 10, 13, 16, 19, 20). The central
stellate cell of the plexus in S. rubiformis is slightly different from that of
S. taiwanensis. The former is a thick-walled structure connecting with a
monohyphal stalk; however, the latter is a swollen cell initiated by fusion
with more than one monohyphal stalk (Figs. 12-15). Among the
interwoven plexal hyphae, flat stellate cells as found in S. rubiformis are
also frequently observed and connected with chlamydospores_ or
Figures 7-11. Micrographs of Sclerocystis spp. Fig. 7. Chlamydospore of
S. coremioides not occluded by a septum. Note a branch (arrow head)
formed on a subtending hypha. Fig. 8. Partial sporocarp of S.
coremioides showing a slender subtending hypha (arrow heads). Fig. 9.
Partial sporocarp of S. liguidambaris showing chlamydospores (S)
embedded in club shaped structures (P) protruding from the central
plexus. Fig. 10. Cross section of sporocarp of S. coremioides showing
chlamydospores arranged radially in a hemispherical layer. Note the
intersporal hyphae connecting between the central plexus and peridium.
Fig. 11. Two naked sporocarps of S. clavispora (SC) and S. taiwanensis.
S. taiwanensis producing remarkable smaller sporocarp. Scale bars in
figures 7, 8, 9 =30 um.
31
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32
intersporal hyphae (Fig. 19). Sporocarps of S. coremioides are
frequently produced in columns or branched structures through lateral or
distal proliferation. Nevertheless, the sporocarps of S. rubiformis are
frequently interconnected into a flat piece (©2 cm diam.) by a
monohyphal stalk. Among the collections of S. taiwanensis, a few young
sporocarps were isolated and all of the chlamydospores were vesicular in
appearance (Fig. 14). The apical wall of chlamydospores is formed last
as sporocarps mature (Figs. 3, 17, 18). A similar type of spore formation
is also found in S. clavispora (Fig. 5).
HYPHAL STALK OF SPOROCARP: One of the main characters in S.
coremioides is its hemispherical sporocarp with a hyphal stipe (Fig. 16).
The hyphal stipe is a rope of interwoven hyphae. However, other species
of Sclerocystis are only associated with one or more monohyphal stalks
(Figs. 12, 14, 20). These monohyphal stalks are usually thick-walled.
Young sporocarps of §. rubiformis with a monohyphal stalk are very
similar to those of S. coremioides, excluding the absence of peridia and
intersporal hyphae (Fig. 12).
GERMINATION OF SPORES: Germinating sporocarps of 5S.
coremioides and S. sinuosa have often been observed. Spores germinate
from intact sporocarps, producing extramatrical vesicle-like structures.
Figures 12-16. Micrographs of Sclerocystis spp. Fig. 12. A young
sporocarp of S. rubiformis showing spores arranged in a broom shape.
Note the plexus connecting with a monohyphal trunk (asterisk). Scale=
30 um. Fig. 13. A fragment of sporocarp of S. sinuosa showing sinuous
peridial hyphae and simple interwoven plexal hyphae. Fig. 14. A
primordial sporocarp of S. faiwanensis connecting with three
monohyphal stalks (arrow heads). Note spores in a vesicular stage (V).
Scale= 30 um. Figs. 15. A swollen central plexus of S. taiwanensis
actively projecting out to form spores. Note the presence of a broad
hyphal trunk (HT) and vesicles (arrows). Scale= 30 um. Fig. 16. Cross
section of sporocarps of S. coremioides showing a hyphal stipe (HS),
central plexus (PL) and spores produced in a hemispherical layer. Central
plexal hyphae are comparatively broader than subtending hyphae of
spores. Scale= 100 um.
33
DISCUSSION
The Species of Sclerocystis included in the present study form a
natural grouping on the basis of method of sporophore formation, spore
germination, chlamydospore and sporocarp. production, ‘and _ the
appearance of the septa and hyphal stalks of the sporocarps (Table 1).
The affinity of S. coremioides is closer to the other species of
Sclerocystis than to Glomus. Although three species of Glomus, i.e., G.
ambisporum, G. heterosporum (Smith and Schenck, 1985), and G.
dimorphicum Boyetchko and Tewari (1986) produce sporocarps similar
to those of S. rubiformis the ability to produce dimorphic
chlamydospores clearly distinguishes them from Sclerocystis. Dimorphic
species, G. ambisporum, G. heterosporum and G. dimorphicum, appear
to be transitional taxa linking G/omus and Sclerocystis, because they
exhibit generic features of both. To date, these three species appeared to
be only distributed in North America. The chlamydospores in sporocarps
of Glomus were mostly arranged randomly and could be enclosed by a
peridium or naked.
Evolutionary trends of sporocarps in Sc/lerocystis are visible
through a comparison of their structures (Fig. 21). Sporocarps of S.
rubiformis are considered as primitive due to their lack of a peridium and
producing sporocarps from a simple, broad, thick-walled plexal cell.
Sporocarp aggregates were not uncommonly isolated and were
interconnected by a single "fecund" hypha which was the same as that
found in G. ambisporum (Smith and Schenck, 1985). In the naked
sporocarps of §. ftaiwanensis , the central plexus is more complicated
than in S. rubiformis and is formed by more than one monohyphal stalk
(Fig. 15). Sclerocystis clavispora produces sporocarps similar to S.
taiwanensis, but the central plexus is replaced by interwoven hyphae.
Nevertheless, broad, thick-walled stellate cells similar to those of
Figures 17-20. Micrographs of Sclerocystis spp. Fig. 17. Crushed young
sporocarp of S. faiwanensis showing thin-walled spores (SP). Scale= 30
um. Fig. 18. Mature sporocarp of S. taiwanensis showing the gradually
thickened apex of spore walls. Note a monohyphal stalk (arrow head)
connecting with plexus. Scale= 30 um. Fig. 19. A broad, thick-walled cell
(arrow head) in the central plexus of a sporocarp of S. liquidambaris.
Scale= 30 um. Fig. 20. Aperidial sporocarp of S. sinuosa showing four
monohyphal stalks (arrow heads). Scale= 100 um.
36
S. rubiformis are often mixed in the plexus. Consequently, S. clavispora
is thought to be advanced S. faiwanensis, and both are senior to S.
rubiformis. Sclerocystis liquidambaris appears to be derived from the S.
clavispora/S. taiwanensis type, with the hyphal branches arising from the
base of the spores becoming thick walled. The peridium of S.
liquidambaris is thought primitive if compared with that of S. sinuosa
and S. coremioides, since it is only tightly packed by thick-walled,
clubshaped cells. In the sporocarps of S. sinuosa and S. coremioides,
however, a peridium of interwoven hyphae is well developed. The
peridial hyphae originate from the central plexus (Figs. 10, 16).
Sclerocystis sinuosa has a number of monohyphal stalks radiating out of
sporocarps. Thus, it resembles S. coremioides, except that the
monohyphal stalks of this species are all combined into a coarse stipe. A
single monohyphal stalk has the ability to produce a broomlike fruiting
body, and further develops into a globose sporocarp as found in S.
rubiformis (Fig. 12). However, the fusing of many monohyphal stalks
into a stipe would result in a lose of the ability to produce spores in a
globular arrangement. The result would be a hemispherical sporocarp.
Results of this study support the retention of the generic concept
of Sclerocystis as proposed by Gerdemann and Trappe (1974). However,
the definition of Sclerocystis is modified as following.
SCLEROCYSTIS Berk. & Br. emend. Wu
Sporocarps globose or subglobose or hemispherical, enclosed by
a peridium or naked, with a multihyphal stipe or monohyphal stalks;
chlamydospores arranged side by side in a single layer, radiating out from
a central plexus of hyphae; central plexus composed of a broad, stellate
thick-walled cell or interwoven hyphae; development of chlamydospores
within sporocarps synchronous or asynchronous; sidebranches frequently
produced from the base of chlamydospores, becoming spores or
intersporal hyphae.
Fig. 21. Diagram of sporocarpic evolution of Sclerocystis spp. A. S.
rubiformis, B. S. clavispora and S. taiwanensis, C. S. liquidambaris, D.
S. sinuosa, E. S. coremioides.
21
OF,
38
TABLE 1. COMPARISON OF SPORE ONTOGENY AND SPOROCARP
FORMATION IN SCLEROCYSTIS
CHARACTERISTICS | S. COREMIOIDES | SCLEROCYSITS spp.
SPOROCARP in chains or branches _| in flat pieces
FORMATION (S. rubiformis)
SPOROCARP vesicle structures vesicle structures
GERMINATION (S. sinuosa)
SPOROCARP STALK monohyphal stalk
(S. rubiformis)
more monohyphal
stalks (S. sinuosa, S.
taiwanensis..etc
SPOROPHORE FROM | present present
BASE OF
CHLAMYDOSPORES
ACKNOWLEDGEMENT
This research was financially supported by National Science
Council of Taiwan, Republic of China (NSC 82-0409-B-055-013).
Appreciation is extended to Drs. R. E. Koske, James W. Kimbrough, and
G. L. Benny for their critical reviews of this manuscript and for providing
specimens used in this research.
LITERATURE CITED
Almeida, R. T. and N. C. Schenck. 1990. A revision of the genus
Sclerocystis (Glomaceae, Glomales). Mycologia 82:703-714.
Sh
Gerdemann, J. W. and J. M. Trappe. 1974. The Endogonaceae in the
Pacific Northwest. Mycologia Memoir No. 5, 76p.
Iqbal, S. H. and B. Perveen. 1980. Some species of Sclerocystis
(Endogonaceae) from Pakistan. Trans. Mycol. Soc. Japan 21:57-
63.
Madelin, M. F. 1979. An appraisal of the taxonomic significance of some
different modes of producing blastic conidia. Pp. 63-80. Jn: The
Whole Fungus. Vol. 1. Ed., B. Kendrick. National Museum of
Canada, Ottawa.
Smith, G. W. and N. C. Schenck. 1985. Two new dimorphic species in
the Edogonaceae: Glomus ambisporum and Glomus heterosporum.
Mycologia 77:566-574.
Trappe, J. M. 1977. Three new Endogonaceae: Glomus constrictus,
Sclerocystis clavispora, and Acaulospora _ scrobiculata.
Mycotaxon 6:359-366.
Wu, C.-G. and Z. C. Chen. 1986. The Endogonaceae of Taiwan. I. A
preliminary investigation on Endogonaceae of bamboo vegetation
at Chi-tou areas, Central Taiwan. Taiwania 31:65-88.
Wu, C.-G. and Z. C. Chen. 1987. The Endogonaceae of Taiwan. I]. Two
new species of Sclerocystis from Taiwan. Trans. Mycol. Soc. Rep.
China 2:73-83.
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Volume XLVI, pp. 41-57 April-June 1993
SYSTEMATIC AND BIOLOGICAL STUDIESIN THE BALANSIEAE
AND RELATED ANAMORPHS. III.
ASCOSPORE AND MACROCONIDIAL GERMINATION
AS A TAXONOMIC CRITERION
RYAN A. PHELPS and G. MORGAN-JONES
Department of Plant Pathology, College of Agriculture and Alabama
Agricultural Experiment Station, Auburn University, Auburn, Alabama 36849
ABSTRACT
Ascospore and Ephelis macroconidial anamorph germination in vitro on
agar media is described in Balansia aristidae Diehl and B. epichloe (Weese)
Diehl, and ascospore germination in Myriogenospora atramentosa (Berk. &
Curt.) Diehl. A review of the literature on the subject of spore germination in
the Balansieae is presented. Consideration is given to the usefulness of
differing germination patterns in the taxonomy of the tribe, including species of
Atkinsonella Diehl, Balansia Speg. and Epichloe (Fr.) Tul. It appears that, in
the taxa so far studied in detail, differences occur that are reflective of both
generic and specific distinctions.
INTRODUCTION
As alluded to in the prologue to this series of papers (Morgan-Jones et
al., 1992), a number of problems exist in the taxonomy of the Balansieae.
These have arisen partly because of a limited number of criteria, including host
relationships, and overall morphology of conidial and perithecial stroma, used in
delimiting taxa. In many instances, species distinctions are not readily apparent
solely on the basis of host specificity and/or stroma morphology. In the case of
Balansia aristidae and B. strangulans (Mont.) Diehl, for example, their
stromata are essentially similar except for the fact that in the former
perithecia are more prominent. Likewise, their Ephelis anamorphs are
virtually indistinguishable. During the course of the present studies, attempts
are being made to evaluate additional criteria of potential taxonomic usefulness.
Among them are details of host-pathogen interactions, in vitro colony growth
characteristics and peculiarities of ascospore and Ephelis macroconidial
germination. It is also anticipated that analysis of molecular sequence data will
aid in resolving questions of relatedness among taxain this complex.
Alabama Agricultural Experiment Station Journal Series Number 18-923366
42
Diehl (1950) reported studying the germination of the Ephelis
macroconidia of six species of Balansia in tap and rain water and on the surface
of sterile agar media. The taxa observed in this regard, except for Balansta
obtecta Diehl, and the composition of the media, were not specified.
Germination of the conidia of Ephelis mexicana Fr., the anamorph of B.
obtecta, was illustrated by a line drawing. No clear distinction was found in
conidial appearance or germination among the species. Both a single primary
apical germ tube and also lateral ones were reported to be produced. Diehl (loc.
cit) described the acicular Ephelis conidia as being characterized by
indentations that were thought to be due to the presence of septa, although these
could not be discerned in most species. A tendency for conidia to break apart at
one or two slight constrictions was also considered to indicate occurrence of
septa. Septa were reported to occur in the macroconidia of Balansia linearis
(Rehm) Diehl and ascospores were said to develop septa before or at
germination. When the latter occurs, each cell may form one or more germ
tubes perpendicular to the axis of the ascospore. Some swelling of the
segments of the ascospores of B. linearis at germination was illustrated.
Ullasa (1969), in a study of Balansia claviceps Speg. in vitro,
investigated conidial and ascospore germination in hanging drop cultures.
Conidia were found to swell considerably in water and water agar prior to
germination. After forty-eight hours, one to five polar and lateral germ tubes
were formed. At this stage conidia became variously septate and individual
cells became somewhat bulbous and produced more than one germ tube. After
seventy-two hours germ tubes branched or remained simple and formed whorls
of secondary conidia at their tips similar in morphology to the parent conidia.
Such conidia either germinated or produced tertiary conidia by a budding
process. On potato dextrose agar slow-growing yeast-like colonies consisting
mainly of masses of conidia were produced. Ascospores germinated similarly
by germ tubes, at the tips of which whorls of conidia were formed.
Luttrell and Bacon (1977) reported that in Myriogenospora atramentosa,
which they classified in the Balansieae, one-celled part-ascospores consistently
germinated by forming an aborted foot at one end and a coiled germ tube at the
other. The germ tube failed to develop into a hypha on common laboratory
media. It was also stated that in species of Balansia each ascospore cell may
germinate by a coiled germ tube produced laterally just below each septum. The
Ephelis anamorph was described as "fusoid conidia produced on phialides at the
surface of the stroma" but no account of conidial germination was given.
Subsequently Rykard et al. (1982) showed the Ephelis state of this species to
have holoblastic, sympodial conidiogenous cells and described conidial
germination. This was found to be characterized by the development of bipolar
or lateral germ tubes. In a liquid medium (referred to as M102), conidia
germinated by forming bulbous, flattened, spoon-shaped ends which produced
elongated germ tubes. On cornmeal malt extract (CMM) and water agar (WA),
branched or unbranched, determinate germ tubes functioned as conidiogenous
cells, producing a sequence of secondary conidia. No mention was made of any
conidial septation.
Conidium production from ascospores, without an intervening hyphal
phase, is known to occur in a number of ascomycetes. Several didymosporous,
43
phragmosporous and dictyosporous species belonging to the Nectria Fr. complex,
for example, produce microconidia, often referred to as ascoconidia, from
ascospores while still remaining within asci. In Nectria aquifolii (Fr.) Berk.
and N. coryli Fuckel numerous microconidia are formed from minute, open-
ended, tube-like extensions at both ends of their one-septate ascospores, and
their ontogeny is considered to be phialidic (Booth, 1959). Likewise,
microconidia are produced from the multiseptate ascospores of Scoleconectria
cucurbitula (Tode:Fr.) C. Booth through minute openings in their walls. (Booth,
loc. cit.). Rossman (1983) described the ascospores of this fungus as ‘budding’
to form ascoconidia. A similar process occurs in species of Thyronectria
Sacc., including T. balsamea (Cooke & Peck) Seeler, T. berolinensis (Sacc.)
Seaver and T. Jamyi (Desm.) Seeler. In all of these taxa ascospores never
produce germ tubes but rather disintegrate following microconidial production.
Ascospore germination in Epichloe typhina (Pers.:Fr.) Tul. was
investigated by Bacon and Hinton (1988) on CMM and water agar. A process,
referred to as iterative germination, was reported to occur in which ascospores
produced lateral, aculeolate, phialidic conidiogenous extensions perpendicular to
their long axis. One such entity was produced from each cell of the ascospore
and sometimes delimited from it by a transverse septum, sometimes not. Each
conidiogenous element produced several conidia terminally. Under varied
conditions, germ tubes leading to hyphal development werenever produced.
Germination of discharged ascospores of Atkinsonella hy poxylon (Peck)
Diehl and A. texensis (Diehl) Leuchtmann & Clay was studied by Leuchtmann and
Clay (1989) on cornmeal-dextrose agar. This was found to be of an identical
pattern in both species. Within the ascus ascospores are usually seven-septate
but split at their median septum, where they are constricted, into two four-
celled part-ascospores prior to germination. Germ tubes were found to be
initially produced, one per cell, at both ends of the part-ascospores and
laterally, immediately below the septa delimiting the end cells. No germ tube
was produced in the vicinity of the median septum of each part-spore. At a
later stage, the part-ascospores often split further, at their median septum, and
additional germ tubes were formed from free ends or from the base of existing
tubes. No study of Ephelis anamorph conidial germination in these species was
reported.
We have had opportunity to study Balansia aristidae, B. epichloe, and
Myriogenospora atramentosa in vivo on grasses in Alabama and the former two
have been brought into pure culture. It is clear from previous reports that
ascospore and/or conidial germination patterns differ between genera and, at
least in some instances, between species of the Balansieae. In order to further
evaluate the possible usefulness of modes of germination in the taxonomy of the
tribe, ascospore and conidial germination characteristics of the two Balansia
species, and ascospore germination in M. atramentosa, have been investigated
and are reported upon herein.
MATERIALS AND METHODS
Plants of Aristida purpurascens Poir., Eragrostis capillaris (L.) Nees,
and Paspalum notatum Fliiggé, bearing stromata of B. aristidae, Balansia
£4
epichloe, and Myriogenospora atramentosa, respectively, were collected from
sites in Lee, Montgomery, and Tallapoosa counties during early to mid summer
1991. Voucher specimens from these collections have been deposited in the
Auburn University Mycological Herbarium (AUA) and at BPI. Infected plants
were transplanted into 15 cm diameter plastic pots containing local sandy loam
and grown in a Conviron growth chamber. An environment of 90% relative
humidity and alternating 12-hour light/dark periods (provided by incandescent
and fluorescent lights) at temperatures of 32° C and 24° C respectively, was
maintained. Using a sterile dissecting needle, macroconidial masses were
scraped from superficial ephelidial pustules on culm-borne stromata of B.
aristidae and from leaf-borne stromata of B. epichloe. No conidia were
harvested from Myriogenospora atramentosa. Ascospores were harvested
from all three taxa by cutting off the tops of several mature, stroma-borne
perithecia horizontally with a razor blade, followed by removal of their exposed
contents with a sterile needle. Both conidia and ascospores harvested by these
methods were immediately suspended in sterile distilled water. Spore
suspensions were then diluted to appropriate concentrations, and plated on each
of three media. The media used during this study were 2% WA, CMM, and EC-
14 (3 g KH>PO,, 2 g K,HPO,-3H20, 2 g NH4S0O,q, 5 g MGSO,-7H30, 40 g
mannitol, 10 g yeast extract, 20 g agar, + water to equal 1 L), a medium
developed by the late Edward M. Clark and Richard A. Shelby at Auburn
University for grass endophyte culture (unpublished). Using a glass rod,
approximately .3 ml of spore suspension was spread onto the surface of the
medium in each petri plate. Ascospores were also harvested by taping mature
perithecial stromata inside the lids of petri plates and awaiting the forcible
discharge of ascus contents onto the agar surface below. After 1, 2, or 4 hours
the stromata were removed and the plates sealed. Germinating spores were
observed in situ with a Zeiss microscope using low power objectives.
Limitations of such observations included compromised resolution and contrast
resulting from the relatively thick agar background, and inability to study the
spores at high magnification due to lack of clearance for an oil immersion lens.
Attempts to transfer germinating spores from petri plates to microscope slides
generally resulted in damage to the spores. To enhance imaging, and enable
viewing at high magnification, spores were allowed to germinate on medium-
coated 22 mm #1 slide coverglasses. Coverglasses were dipped in hot agar
medium and placed inside petri plates on the surface of the same medium to
prevent desiccation. After adding the spore suspension to the upper exposed
surface of the coverglasses and allowing the excess liquid to evaporate, the
petri plates were sealed and incubated at 25° C. Inoculated coverglasses were
PLATE 1. Balansia aristidae. A-F, Ephelis conidia; A, from nature; B, initial
swelling on agar medium; C, apical swelling (early stage); D, apical swelling
(later stage); E & F, apical swelling (final stages), and initiation of primary
germ tubes: G-L, part-ascospores; G & H, from mature perithecium (median
septa indicated by arrows); I, median disarticulation and terminal swelling
(early stage); J-L, median disarticulation and terminal swelling (later stages),
lateral germ tube initiation (indicated by arrowheads) and disarticulation of
swollen ends.
45
46
removed periodically during germination, inverted, and mounted on hanging-drop
slides with the surface bearing spores downward. Prior to viewing, excess
agar was gently wiped from the upper surface of the mounted coverglass.
When standard slides were used instead of hanging drop slides, germinating
spores were frequently damaged.
RESULTS
Balansia aristidae. Conidia.
Conidia of this species are long, cylindrical to somewhat fusiform,
obtuse at the apex, subtruncate at the base, 10 to 15 ym long and less than 1 pm
wide (Plate 1, A; Figure 1, A). Germination by germ tube occurred after 4
days on EC-14 and CMM media and was preceded by morphological changes.
Following 48 hours on these media, conidia swelled appreciably at their apex to
form an ellipsoid to oblong portion about 3 ym in length and 2 ym wide, while the
opposite, basal end remained more or less truncate (Plate 1, A & B; Figure 1,
B). After 5 days the swollen portion became much more elongate (up to 7 pm),
assuming a long-oblong or more or less cylindrical aspect or, in some cases,
became narrowly obpyriform (Plate 1, D; Figure, 1, B). Initial germination
was mostly monopolar but variable. A primary germ tube often originated from
the conidium immediately below the base of the terminal swelling or a short
distance below it (Plate 1, E & F). Alternatively, more rarely, a germ tube
originated apically. Occasionally this was followed shortly thereafter by
disarticulation of the terminal swelling from the remainder of the conidium.
After 10 days, some swollen terminals became up to 12 pm long where apical
germination did not occur. Following initial germ tube production a secondary
germ tube sometimes originated proximal to the first. In some instances,
following a period of 12 days, a germ tube was also produced from the opposite,
basal end of the conidium. Many conidia underwent the described morphological
changes but failed to germinate after 10 days.
Although most conidia failed to germinate on WA, germination followed a
similar course except that terminal swelling was delayed 24 to 48 hours and
never developed beyond 3.5 ym in length. After 12 days a small percentage
developed a swollen spherical structure up to 6 pm in diameter (Figure 1, C),
located distal to the already swollen oblong, terminal portion, frequently giving
rise to one or two germ tubes. Such germ tubes generally developed into
extensive hyphal systems. Though rare, hyphae sometimes also emerged from
the basal end.
PLATE 2. A, Balansia aristidae part-ascospores 72 hours after direct
discharge onto agar medium; B-G, part-spores after 4 to 7 days on agar media
showing various stages of terminal swelling, median and terminal septation and
disarticulation (indicated by arrowheads), and sympodial germ tubes; H,
chlamydospore-like cells arising from a part-spore on agar medium; I-L,
Myriogenospora atramentosa part-ascospores showing stages of germination
and germ-tube branching.
47
48
C
(Bike Be a et}
Sym
FIGURE 1. Balansia aristidae, Ephelis conidia. A, from nature; B & C, stages
and patterns of germination.
Spm |
FIGURE 2. Balansia aristidae part-ascospores. A, from nature; B & C, various
stages of germination, septation and disarticulation.
50
Ascospores.
Ascospores of Balansia aristidae disarticulate within the asci, forming
one-septate part-spores which are then discharged. Part-spores are narrow
and cylindrical with truncate ends, 21 to 34 jm in length and less than 1 pm wide
(Plate 1, G & H; Figure 2, A). After 48 hours on EC-14 medium, part-spores
swelled at both ends forming bulbous, ellipsoid terminals 3.5 to 4.5 pm in length
and disarticulated at the median septum (Plate 1,1; Figure 2, B). Following
another 48 hours, the swollen ends became nearly 9 pm long. Lateral germ
tubes developed directly or a short distance below the terminal swellings (Plate
1, H; Plate 2, B& C). Concurrently or immediately before germ tube initiation,
a septum was laid down just above its point of origin. Disarticulation of the
swollen terminal from the remainder of the half part-spore frequently occurred
at this septum (Plate 2, D). After 7 or more days, some swollen ends reached
14 zm in length and a second germ tube was occasionally produced from the base
of the half part-spore (Plate 2, G; Figure 2, C). A few half part-spores
swelled appreciably, becoming multiseptate and forming thick-walled,
melanized chlamydospore-like structures (Plate 2, H).
Part-spores behaved similarly on WA. Septae and terminal swellings,
however, were not as well-defined, and spores became generally more
irregular in shape. Lateral and terminal germ tubes occurred frequently after
48 hours, and quickly developed into mycelia. Thick-walled, chlamydospore-
like cells also began to occur at 48 hours. After 4 days these structures were
extremely common. Following 7 days, part-spores had either failed to swell
and began deteriorating, formed mycelia, or formed chlamydospore-like
structures.
Balansia epichloe. Conidia.
Conidia of this species are acicular to fusiform, acute at the apex,
subtruncate at the base, 16 to 21 um long and less than 1 pm wide (Plate 3, A;
Figure 3, A). Within 24 hours on EC-14 medium conidia swelled slightly and
germinated by apical germ tubes, one per conidium. Moderate swelling occurred
evenly over the basal two-thirds of the spore, the apical portion remaining more
attenuate (Plate 3, B). Just prior to apical germination, the extreme tip swelled
slightly (Plate 3, C). After 24 hours germ tubes averaged 7 pm, emerging from
a Slight enlargement at the apex of the tapered conidium (Figure 3, B). A
similar enlargement developed at the basal end also. After 48 hours apical germ
tubes approached 12 pm in length. Germination remained monopolar for nearly
PLATE 3. Balansia epichloe. A-H, Ephelis conidia; A, from nature; B, initial
swelling; C & D, further swelling and early stage of germination; E, well-
developed apical germ tubes; F & G, disarticulation of initial germ
tube(indicated by arrowhead) and initiation of sympodial germ tube; H, bipolar
germination; I-L, part-ascospores; I, from mature perithecium; J & K, median
disarticulation (indicated by arrowhead) and terminal germination; L, sympodial
development of secondary germ tubes (indicated by arrows) and disarticulation
of initial terminal germ tubes (indicated by arrowhead).
Hy
oy
B
)
FIGURE 3. Balansia epichloe, Ephelis conidia. A, from nature; B & C, stages
and patterns of germination.
A
eee
>
a
Fagen TE Ome a
FIGURE 4. Balansia epichloe part-ascospores. A, from nature; B, various
stages of germination.
54
5 days, at which time a second germ tube occasionally developed from the basal
enlargement (Plate 3, H). More commonly, sympodial initiation of a second
germ tube occurred at the apical end of the spore (Plate 3, F & G). Subsequent
disarticulation of the initial germ tube typically ensued.
Germination rarely occurred on WA, and only after 72 hours. The
characteristic swelling prior to germination was more pronounced than on EC-
14 medium. Following 5 days on WA germ tubes up to 5.5m had developed.
Though infrequent, some conidia swelled appreciably, approaching 40 pm in
length and 2.5 ym in basal width, the apical half remaining quite narrow. Even
after 10 days germination failed to progress further.
Ascospores.
Ascospores of Balansia epichloe disarticulate within the asci, forming
one-septate part-spores which are then discharged. Part-spores are broadly
filiform with truncate ends, 32 to 46 ym in length (Plate 3, I; Figure 4, A).
Although half part-spores seldom separated completely, their partial
disarticulation at median septae was a common phenomenon. Germination on
EC-14 medium occurred within 24 hours. A germ tube developed from the outer
exposed end of each half part-spore, and usually grew sharply to one side or the
other (Plate 3, J &K). Since half part-spores typically remained connected,
this germination pattern resembled bipolar germination of the part-spore. After
72 hours germ tubes had reached at least 10 to 14 um in length on WA, and at
least 17 pm in length on EC-14. No further development occurred on WA. On
EC-14 medium, sympodial initiation of secondary germ tubes, and disarticulation
of initial germ tubes followed (Plate 3, L; Figure 4, B).
Myriogenospora atramentosa. Ascospores.
The germination pattern characteristic of ascospores of this species is
particularly distinctive. Long, narrow, filiform, aseptate part-spore with
acutely tapered ends are discharged from each ascus. Rarely, these part-spores
remain joined end-to-end in pairs. Within 48 hours on WA and EC-14 media,
part-spores exhibited monopolar germination by the development of a germ tube
from a slight apical enlargement. A similar swelling on the other end remained
dormant throughout germination, which continued for several days and then
ceased. At first germ tubes were distinctly curled, becoming undulate or
loosely circinate to somewhat coiled (Plate 2, I-L; Figure 5, B). On EC-14
medium, branching was frequent, each branch continuing the same growth habit.
FIGURE 5. A, Myriogenospora atramentosa part-ascospores from nature; B,
part-spores germinating; C, germinating ascospores of Atkinsonella hy poxylon
(after Leuchtmann and Clay, 1989); D, segment of Epichloe typhina ascospore
showing iterative germination (after Bacon and Hinton, 1988).
56
DISCUSSION
Ascospore and conidial germination peculiarities clearly vary appreciably
between the member genera of the Balansieae and, to a lesser extent, between
species. It is interesting to note that ascospore germination in the two species
of Balansia investigated during the course of this study is more closely similar
than between these and species of Atkinsonella. Rykard et al. (1984) questioned
the desirability of maintaining Atkinsonella and Balansiopsis Hohnel as separate
genera from Balansia. Further information on ascospore germination in
additional species of Balansia will, perhaps, help resolve that question. As
pointed out by Leuchtmann and Clay (1989), disarticulation of seven-septate
ascospores into two three-septate part-spores, and the number and location of
germ tubes formed subsequently, appears to be unique for the genus
Atkinsonella. This is certainly different from species of Balansia where the
released part-spores are one-septate and those of Myriogenospora which are
aseptate. That the two known species of this genus, A. hypoxylon and A.
texensis, have identical ascospore germination (Leuchtmann and Clay, loc. cit.)
indicates that they are closely related. The latter, on morphological grounds
alone, was originally classified as A. hypoxylon var. texensis by Diehl (1950).
It was recently recognized as a separate species mainly on the basis of host
association, isozyme variation, colony differences in vitro and host cross-
inoculation incompatibilities (Leuchtmann and Clay, loc. cit.).
The distinctive ascospore discharge and germination characteristics of
Epichloe typhina and Myriogenospora atramentosa are further indicative of the
usefulness of this feature in determining relatedness. The single-celled part-
spores of the latter and the very different germination morphology are
reflective of the genetic distance between Myriogenospora, a monotypic genus,
and others classified in the tribe Balansieae. The monopolar germination
characteristic of each part-spore also indicates this. Myriogenospora was not
included by Diehl (1950) in the tribe but added by Luttrell and Bacon (1977)
despite its considerably different epiphytic habit and ascus morphology. Luttrell
and Bacon (loc. cit.) considered the type of germination in M. atramentosa to be
significant, noting that in species of Balansia each cell in the filamentous
ascospore may germinate by a coiled germ tube produced from the side of the
cell just below the septum. That is, of course, a simplification and
generalization for, as indicated in the results noted above, there are variable
patterns of ascospore disarticulation, secondary septation, and secondary
breaking apart of part-spores, associated with the sequence of germ tube
initiation. Germ tubes in Balansia do not usually appear to be undulate, circinate
or coiled in the manner of those formed from ascospores of Myriogenospora.
The swelling of the Ephelis conidia of M. atramentosa in liquid culture
reported and illustrated by Rykard et al. (1982) is, interestingly, reminiscent of
the morphological changes, described herein, undergone by conidia of Balansia
aristidae prior to germination. The latter differs, however, in that the
swelling occurs at the apex of the conidium only and not at each end, as was
noted to occur in M. atramentosa by Rykard et al. (loc. cit.). The sequence of
secondary conidia sympodially produced from the tip of a determinate germ tube
in M. atramentosa form in essentially the same manner as the primary conidia.
57,
Such sympodial growth is also seen occurring during germination of conidia and
ascospores of both B. aristidae and B. epichloe.
The sequences of events occurring during macroconidial and ascospore
germination in Balansia aristidae and B. epichloe are distinctive for each
species and, therefore, of use in their characterization. The considerable
degree of difference in ascospore growth pattern, morphology and
disarticulation is of particular interest. The secondary septation and the
eventual breaking up of the original one-septate part-spore into four separate
cells in B. aristidae is, to our knowledge, unique in the genus. There is, in this
case, a sequence of primary, secondary and tertiary disarticulation of the
original ascospores delimited within the asci. Whether such differences exist
between other species of the genus remains to be determined. The account by
Ullasa (1969) of germination in Balansia claviceps in hanging drop cultures is
not comparable to the present study because of the differing conditions and less
detailed observations. It would be interesting, for example, to investigate if
there are differences in regard to germination between B. aristidae and B.
strangulans, two species that are considered close (Diehl, 1950), and likewise
between B. epichloe and B. henningsiana (Moell.) Diehl, two species that are
superficially similar, and from time to time have been considered to be the
same fungus, but are now recognized as separate and distinct entities. There is
reason to believe that, even if all individual species lack clear-cut differences
in respect to germination peculiarities, this criterion is nevertheless of some
use in indicating relatedness and/or separateness.
ACKNOWLEDGEMENT
This research was supported by a National Science Foundation (BSR-
8922157) grant to J. F. White, Jr. and G. Morgan-Jones. We thank Dr. J.
Leland Crane for reviewing this paper.
LITERATURE CITED
BACON, C.W. and D.M. HINTON. 1988. Ascosporic iterative germination in
Epichloe typhina. Trans. Br. mycol. Soc. 90: 563-569.
BOOTH, C. 1959. Studies of Pyrenomycetes. IV. Nectria (part 1). Mycol.
Pap. 73: 1-115.
DIEHL, W.W. 1950. Balansia and Balansiae in America. Agric. Monograph No.
4, USDA, Washington, D.C. 82 pp.
LEUCHTMANN, A. and K. CLAY. 1989. Morphological, cultural and mating
studies in Atkinsonella, including A. texensis. Mycologia 81: 692-701.
LUT TRELL, E.S. and C.W. BACON. 1977. Classification of Myriogenospora
in the Clavicipitaceae. Can. J. Bot. 55: 2090-2097.
MORGAN-JONES, G., R.A. PHELPS and J.F. WHITE, JR. 1992. Systematic
and eee studies in the Balansieae. I. Prologue. Mycotaxon 43:
401-415.
ROSSMAN, A.Y. 1983. The phragmosporous species of Nectria and related
genera. Mycol. Pap. 150: 1-164.
RYKARD, D.M., E.S. LUT TRELL and C.W. BACON. 1982. Development of the
conidial state of Myriogenospora atramentosa. Mycologia 74: 648-654.
eee on 1969. Balansia claviceps in artificial culture. Mycologia 61:
72-579.
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Volume XLVI, pp. 59-65 April-June 1993
Chaetospermum chaetosporum (Coelomycetes). First record
from the Iberian Peninsula
M. Muntafiola— Cvetkovic* and A. Gémez-— Bolea*
*Department of Plant Biology, Unit of Botany, Faculty of Biology, University
of Barcelona, Diagonal 645, 08028 Barcelona, Spain.
Abstract. Chaetospermum chaetosporum (Pat.) Smith & Ramsb.
was encountered on graminicolous leaves and sheaths with small
distorted areas, in the Tibidabo mountain, Catalonia. The
conidiomatal structures of the fungus showed an_ evident
resemblance to gelatinous sporodochia. This is the first record of
the above mentioned taxa in the Iberian Peninsula.
Key words: Coelomycetes, Chaetospermum, C. chaetosporum.
In July 1990 several graminicolous decaying plants were collected in the
Tibidabo mountain, near the city of Barcelona (Catalonia, Spain), showing
coloured, slightly thickened, distorted areas on the leaves and sheaths. A
closer examination of the lesions revealed the presence of orange —yellow
structures, gelatinous to the touch, resembling groups of sporodochia. The
microscopic preparations made from such crushed structures showed abundant
subcylindric conidia with bipolar appendages characteristic of the genus
Chaetospermum Sacc. The sporodochium —like bodies and the Chaetospermum
conidia were seen in subsequent collections of similar lesions on leaves of
decaying Gramineae growing near the same place. Some conidia of other
fungi (mainly Alternaria and Cladosporium species), together with those of
Chaetospermum, were observed eventually in the slides, as well as some few
conidia of Drechslera (probably D. victoriae (Meehan & Murphy) Subramanian
& Jai).
Since no published reports have been found concerning the presence of
Chaetospermum species in the Iberian Peninsula, the description of the fungus
encountered in the Tibidabo is herein presented.
60
Material and Methods
Collections studied. The material studied consisted of decaying grass
leaves and sheaths showing small distorted areas with minute sporodochium —
like conidiomata. The material was collected in the Tibidabo mountain, Serra
de Collserola (S00 m altitude), near Barcelona, Catalonia, Spain, the 1st, 7th
and 8th July 1990, a month with no rains, that followed a dry period.
Microscopic preparations and examination. The conidiomata were removed
from the plant substrates with sterilized needles and mounted onto glass
microscope slides in lactophenol slightly stained with safranin. Preparations
fro, thirteen different collections were sealed with a nailpolish double coat
and labeled EE. 90, 30-42. Microscopic examinations were made using a
Leitz Diaplan microscope and 1,000 magnifications by oil—immersion, bright
field, phase-contrast and the Nomarski interference optic systems.
Micrographs were taken with a Nikon F-—610 camera and Ilford Pan F, 50
ISO film.
Slides labeled EE. 90. 30 and 33 were deposited in the Mycological
Herbarium BCC (University of Barcelona).
Results
The morphologic features of the fungus encontered in the Tibidabo
mountain were the following:
Conidiomata consisting of orange—yellow, sessile, gelatinous, mostly
aggregate and roughly spherical structures more similar to sporodochia than
to typical acervula. Conidiophores constituted by tufts of hyaline hyphae,
mostly 30-40 x 15-2 wm (Fig. 1, arrowed), with terminal or lateral
conidiogenous cells, swelling at their tips to give rise the conidia. Conidia
elongate — cylindrical, straight to slightly bent smooth—walled, hyaline, (15—)
30-35 (-45) x (5-) 7.5 (-10) pm, aseptate, with granular contents
(Figs. 1 and 2), with the apices irregularly obtuse or sometimes almost
truncate and bearing three to six terminal or subterminal appendages.
Appendages filiform, flexuous, not branched, not septate, 1 wm _ width,
generally as long as the conidium body, or slightly shorter; with no
protoplasmatic continuity with the conidium body when the internal contents
retracts.
Figs. 1 and 2. Chaetospermum chaetosporum (Pat.) Smith & Ramsb. (Mycol. BCC
Herbarium NO EE. 90. 30). Conidiophores (arrowed) and conidia showing the granular
protoplasm conspicuously retracted (separated) from the cell—wall, and the apical and
subapical insertion of the appendages. Scale bars = 10 um.
61
62
Discussion
The several generic names that have been given to the fungi now
gathered in the genus Chaetospermum reflect the views of their authors
concerning the conidiomatal structure of these fungi. The conidiomata,
initially roughly spherical and later rupturing irregularly, show such,a variation
in morphology that the species have been placed in the Tuberculariales,
Melanconiales and Sphaeropsidales. Patouillard (1888) considered a
sporodochium the conidiomata of the fungus that he described as
Tubercularia chaetospora, for which Saccardo (1892) erected the genus
Chaetospermum, with the following short diagnosis: Sporodochia gelatinosa.
Sporophora ramosa. Conidia ovoidea, hyalina, utrinque 7-8 setosa. Later on
the conidiomata of Chaetospermum were again referred to as sporodochia
(Smith & Rambsbottom, 1913; Agnihothrudu, 1962). However, the
Chaetospermum conidioma was considered essentially a pycnidium by other
authors (Héhnel, 1924; de Fonseka, 1960; Nag Raj & Kendrick, 1972), while
after Sutton (1980) it is acervular. The conidiomatal structures of the fungus
collected in the Tibidabo evidently grade into sporodochial conidiomata.
Sutton (1980) put into synonymy with Chaetospermum the genera
Ciliospora Zimmerman, Mastigonema Speg., Entomopatella Petrak, and
Chaetospermella Naumov, as he had discussed early (Sutton, 1977). Of the
five Chaetospermum species keyed by Agnihothrudu (1962), Sutton (1980)
tentatively recognized only three, mainly based on conidium size. This author
maintained C. gelatinosum Petch in the genus, although recognized that the
conidia are much thicker—walled than in the other accepted taxa and the
appendages are very thin, perhaps not "cellular" in origin.
In Tab. 1 the main morphometric characters of the fungus collected in
the Tibidabo have been compared with those of C. chaetosporum, the type
species of the genus, and other congeneric taxa. It can be concluded that the
Chaetospermum structures found on the plants collected in the Tibidabo fit
well with the morphology and dimensions of C. chaetosporum. It could be
only remarked here that septate spores like those reported and illustrated by
de Fonseka (1960) from 3-4 month-old conidiomata were not seen in our
collections, although the retraction of the conidium body protoplasm observed
in some slides could indicate that the fructifications were quite old. The
retraction was not considered a manipulation artefact, since the phenomenon
was observed in water preparations as well as in the lactophenol — mounted
slides.
Much attention was paid in this study to the appendages of the conidia
since many species of the Coelomycetes show similar structures. Their
features have been considered by Sutton & Sellar (1966) and Sutton (1968,
1973) of great importance. Sutton’s criteria have been sustained by Pirozinski
& Shoemaker (1971), Nag Raj (1981, 1988) and others. According to these
criteria the conidial appendages of the fungus found in the Tibidabo must be
considered of "cellular" origin.
63
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64
C. chaetospermum appears not to be a common fungus in the Iberian
Peninsula. Records of this species or other related taxa are not found in the
literature. Since July 1990 we were not able to collect again this species.
C. chaetosporum has been reported on stems, twigs and leaves of a
variety of plants belonging to very different genera, but no studies have been
found emphasizing this organism as a pathogenic one. However, the lesions
observed on the leaves and sheaths from which the fungus was recovered in
the present study were similar to those attributed by Sprague (1950) to
Dilophospora alopecuri (Fr.) Fr. Controlled inoculation tests of healthy plants
with C. chaetosporum should dilucidate the questions related to _ its
pathogenicity, specially taking into account that conidia of other fungi were
found in the samples examined.
Acknowlegments
M. M.-C. is grateful to the staffs of the "Centre de Documentacié i
Experimentacié de Ciéncies", Barcelona, and of the "Departament de Biologia
Vegetal, Unitat de Botanica, Facultat de Biologia", Barcelona, for the use of
their facilities.
The authors are indebted to Profs. X. Llimona and J. Guarro for
critically reading the typescript.
References
AGNIHOTHRUDU, V. 1962. Notes on fungi from North-East India X.
Chaetospermum camelliae sp. nov. on tea (Camellia sinensis (L.) O.
Kuntze). Mycopathologia et Mycologia Applicata 16, 113 — 116.
FONSEKA, R.N. de 1960. The morphology of Chaetospermum chaetosporum.
Transactions of the British mycological Society 43, 631-636.
HOHNEL, F. von 1924. Uber die Gattung Chaetospermum Sacc. Mitt. bot.
Lab. techn. Hochsch. Wien, 3, 86 — 88.
NAG RAJ, T.R. 1981. Coelomycete systematics. In The biology of conidial
fungi. Vol. I. Edited by G.T. Cole & B. Kendrick. Academic Press: New
York.
NAG RAJ, T.R. 1988. Genera coelomycetarum. XXIV. Chithramia anamorph
gen. nov. Canadian Journal of Botany 66, 903 — 906.
65
NAG RAJ, T.R. & KENDRICK, W.B. 1972. Genera Coelomycetarum III.
Pestalozziella. Canadian Journal of Botany 50, 607-617.
PATOUILLARD, M.N. 1888. Note sur une Tuberculariee graminicole.
Bulletin de la Société Mycologique de France 4, 39-40.
PIROZYNSKI, K.A. & SHOEMAKER, R.A. 1971. Some Coelomycetes with
appendaged conidia. Canadian Journal of Botany 49, 529-541.
SACCARDO, P.A. 1892. Sylloge Fungorum 10 (p. 706). Padua.
SMITH, A.L. & RAMSBOTTOM, J. 1913. New or rare fungi. Transactions
of the British mycological Society 4, 318-330 (p. 328).
SPRAGUE, R. 1950. Diseases of cereals and grasses in North America
(Fungi, except smuts and rusts). Ronald Press: New York.
SUTTON, B.C. 1968. Kellermania and its generic segregates. Canadian
Journal of Botany 46, 181-196.
SUTTON, B.C. 1973. Coelomycetes. In the Fungi. An Advanced Treatise.
Vol. IV A. Edited by G.C. Ainsworth, F.K. Sparrow & A.S. Sussman.
Academic Press: New York.
SUTTON, B.C. 1977. Coelomycetes VI. Nomenclature of generic names
proposed for Coelomycetes. Mycological Papers 141, 1-253.
SUTTON, B.C. 1980. The Coelomycetes. Fungi Imperfecti with Pycnidia
Acervuli and Stromata. Commonwealth Mycological Institute: England.
SUTTON, B.C. & SELLAR, P.W. 1966. Toxosporiopsis n. gen., an unusual
member of the Melanconiales. Canadian Journal of Botany 44, 1505 -
1513;
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MY COTAXON
Volume XLVI, pp. 67-69 April-June 1993
LECTOTYPIFICATION OF OPHIOBOLUS TRICHELLUS
(DOTHIDEALES, ASCOMYCETES)
Christian Scheuer
Institut fiir Botanik der Karl—Franzens—Universitat Graz,
Holteigasse 6, A—8010 Graz, Austria
A lectotype specimen is designated for Ophiobolus trichellus SACC., BOMM. &
ROUSS. The species, recently placed in Tubeufia by SCHEUER (1991), is briefly
described and illustzated.
Key words: Tubeufia trichella, taxonomy.
SCHEUER (1991: 814) has recently published a new combination of Ophiobolus
trichellus SACC., BOMM. & ROUSS., Tubeufia trichella. Although the British material
examined agrees with the original description in every respect, the new combination was
somewhat doubtful, because no type material of Ophiobolus trichellus had been
available at that time. A quite satisfactory syntype specimen from Knocke (Belgium)
could be located in PAD. It agrees with the protologue and with the British material
studied and illustrated by SCHEUER (1991). No specimen from Oostende (Belgium),
the second locality given by the authors in the protologue (in BOMMER & ROUSSEAU
1890: 259), could so far be found in any herbarium. The syntype specimen in BR (Sur les
feuilles d'dmmophila arenaria, [Belgium,] Knocke, Aoit 1890, leg. M. ROUSSEAU)
consists only of a small leaf fragment bearing no ascomata of Ophiobolus trichellus.
Nevertheless, because of the two localities given in the protologue, a lectotype must be
designated.
Ophiobolus trichellus P. A. SACCARDO, BOMMER & ROUSSEAU
in BOMMER & ROUSSEAU, Bull. soc. roy. bot. Belgique 29: 259 (1890),
later cited erroneously as 'O. trichellus BOMM., ROUSS. & SACC.' in Syll.
Fung. 9: 934 (1891).
68
= Ophiochaeta trichella (SACC., BOMM. & ROUSS.) SACC.,
Syll. Fung. 11: 352 (1895).
= Tubeufia trichella (SACC., BOMM. & ROUSS.) SCHEUER,
Mycol. Res. 95(7): 815 (1991).
Ascomata (Pseudoperithecia) + scattered on dead leaves of Ammophila arenaria (L.)
LINK, usually epiphyllous, nesting among the leaf hairs, brown, + globose, ca. 100 um
in diam., with a fringe of dark brown, simple setae around the ostiole. No branched setae
have been found in the lectotype material. Setae ca. 20-40 um in length, tapering from
the base, but always rather torulose and irregular in outline, thick-walled, usually non-
septate. The ostiole is surrounded by small, knob-like peridial cells. Peridium in surface
view composed of irregular, angular cells up to ca. 8 wm in diam., consisting of 3 - 5
layers of slightly flattened cells. Pseudoparaphyses present in mature loculi, ca 1.5-2 um
thick. Asci bitunicate, + cylindrical, 45-66 x 10-15 “~m (SACCARDO, BOMMER &
ROUSSEAU 1890), 8-spored. Ascospores hyaline, flexuous, cylindric-filiform, ca. 48-
65 x 2.5 wm, 14—-16-(?18-)septate, guttulate. The ascospores are usually coiled in the
ascus, before the ascus expands at maturity.
LECTOTYPUS designated here: A la face inférieure des feuilles du Psamma arenaria
[=Ammophila arenaria], [Belgium,] Knocke; ex Herb. E. BOMMER & M. ROUSSEAU
(Herbarium Mycologicum P. A. SACCARDO, 2729, PAD).
Lectotype of Ophiobolus trichellus: (A) Ascoma. (B) Ascospores (lactophenol
permanent slide). — Bars = 20 um.
69
This material differs only very slightly from the British collections studied and
illustrated by SCHEUER (1991: 814, 815). No branched setae have been found in the
lectotype, but in some (not all!) of the British collections. In addition, apparently some of
the ascospores of the two ascomata mounted from the lectotype have more septa (up to
18?) than those of British material (up to 15).
Thanks to Dr. Sergio CHIESA (Padova) for his help in tracing the type specimen in
PAD, and to PD. Dr. Orlando PETRINI (ETH Ziirich) for acting as pre—submission
reviewer.
References
BOMMER, E. & M. ROUSSEAU 1890. Contributions a la flore mycologique de
Belgique. — Bulletin de la societé royale de botanique de Belgique 29: 205-302.
SCHEUER, Ch. 1991. Taphrophila (Dothideales: Tubeufiaceae) and two new species
of Tubeufia with dark setae. - Mycological Research 95(7): 811-816.
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MYCOTAXON
Volume XLVI, pp. 71-80 April-June 1993
TAXOMYCES ANDREANAE, A PROPOSED NEW TAXON
FOR A BULBILLIFEROUS HYPHOMYCETE ASSOCIATED
WITH PACIFIC YEW (TAXUS BREVIFOLIA)
GARY STROBEL and ANDREA STIERLE
Department of Plant Pathology
Montana State University
Bozeman, Montana 59717
DON STIERLE
Department of Chemistry
Montana College of Mining Technology
Butte, Montana 59701
and
W. M. HESS
Department of Botany & Range Science
Brigham Young University
Provo, Utah 84601
ABSTRACT
Taxomyces andreanae is described as a novel
endophytic fungus associated with the inner bark
of Taxus brevifolia Nutt. (Pacific yew). This fungus
has small septate hyphae which average 1.2 wm and
large septate hyphae which average 3.75 wm in
diameter. Hyphal cells of this fungus are
multinucleate. It characteristically forms clumps
of loosely constructed cells (bulbil-like). These
clumps are of various shapes and sizes typically
ranging from ca. 5x5 to 16x30 wm in diameter and
length. The nucleated cells in these clumps
average about 1.5x2.5 wm and appear to be loosely
packed in the bulbil and are incapable of
germination. This fungus grows rapidly on many
common laboratory media, covering the plates with
its mycelium in 3-4 days. itaelacks cclamp
connections, and dolipore septations. Its
telemorph is unknown.
72
INTRODUCTION
The Pacific yew (Taxus brevifolia Nutt.) is the primary
source of taxol, a promising anticancer drug isolated from
the inner bark of this tree (1,2). In a search for microbes
that may influence the production and/or fate of taxol (and
related taxanes)in this tree, we have made a concerted effort
to isolate and identify parasitic and endophytic microbes
associated with it. In the present report we describe a
novel endophytic hyphomycete, Taxomyces andreanae, that was
isolated exclusively from the inner bark on small limbs of a
specific yew tree in Northern Montana.
MATERIALS AND METHODS
The fungus was isolated from the upper limbs of a shrub-
like Pacific yew tree growing as undercover in a mature,
undisturbed cedar forest in Flathead County, Montana. Small
limbs (0.5 - 1.0 cm) were surface treated with 70% ethanol.
The outer bark was peeled back and pieces of the white inner
bark (phloem/cambium) were aseptically removed and placed on
H,O agar. Hyphal tips of fungi growing from the pieces of
the plant were placed on mycological agar and fungal growth
was enhanced. Subsequent attempts to isolate the fungus from
other sites in Montana, Idaho, Oregon and Washington were
unsuccessful.
The growth pattern of the fungus was studied on other
additional plant species. Leaf, stem, and bark samples of
various plant species growing near the tree from which this
fungus was obtained were collected near the Hungry Horse dam
site on the Flathead National Forest. Small pieces (0.5 - 2
cm) of leaves, stems and bark of these species were placed
over 4 layers of cheesecloth, thoroughly dampened, and
autoclaved. Agar blocks (1.0x1.0 cm) supporting fungal
growth were then placed on the sterilized plant material and
fungal growth observed and measured after 1 week.
The growth of the fungus was measured daily after
placement of 0.5x0.5 cm agar blocks on standard freshly
prepared agar media plates (Difco) eg. potato dextrose broth
agar, nutrient agar, oatmeal agar, cornmeal agar, lima bean
agar, water agar, and malt agar.
Agar blocks having mycelia and bulbil-like structures
were fixed and dehydrated as for transmission and scanning
electron microscopy (SEM) (3,4,5,6). For SEM, the material
was then critical point dried, gold coated and sputter
coated, and observed with a JEOL 840A scanning electron
microscope. Fungal structures were measured on SEM
micrographs after critical point drying of tissues. This
1S,
drying procedure causes some shrinkage of biological
structures (ca. 10%) which means that they are probably
Slightly larger, and the clumps of cells more tightly packed
when in the living state.
Nuclei were stained with the 4,6 =diamidino-2-
phenylindole reagent (5ug/ml of 20% ethanol) and observed
under UV at 365 nn.
RESULTS AND DISCUSSION
Taxonomic Treatment and Description
Taxomyces andreanae Strobel, Stierle and Hess gen. et sp.
NOVen 4 ELOS s.1 =6)i.
Fungus endophyticus e cortice interiora Taxo brevifolo
Nutt.; hyphae dimporphae -- parvae 1.25 wm et magnae ca
3.75 um latae et longae; bulbilus cellularum ca 1.25x2.5 um
et laxe contiguus et apparenter non germinans; mycelium
celiter crescens, hyphis fibulis nullis et doliporis septis
nullis; telemorphus ignotus.
Mycelium superficial, composed of a network of highly
branched, septate, multinucleate, usually hyaline, smooth
walled hyphae. Smaller hyphal cells average 1.25 pum in
diameter. Larger cells average 3.75 wm in diameter (Figs. 1-
2). Cells are budded from fructigenous hyphae forming clumps
which vary enormously in shape from spherical to ovoid to
longiform and in size from 5x5 wm in diameter (for spherical
bulbils) to 16 - 30 wm in length (for elongate bulbils).
Bulbil cells remaining colorless. The cells seem to be
loosely packed in the bulbil and are ovoid ca. 1.5x2.5 wm and
are never observed to germinate.
The "clumps" of cells in Fig. 2 and 3 are not located on
sterigmata, but seem to arise by a "budding process" from
hyphal pores (Fig. 3). The number of cells in each clump
Variesawidely (Pig. £2 &) 3)". We have referred to these
"clumps" as bulbils after the broad definition of deBary (7),
thatsers, "Small pluricellular bodies incapable of
germination". In 7. andreanae, the cells of the bulbils, unlike
most bulbilliferous fungi, appear to be loosely packed, but
nevertheless connected with fibrous material (Figs. 3,6).
These clumps of cells might also be considered as conidial
masses but since germination has never been observed (in
sterile H,O and nutrient broth), the clumps of cells seem to
better fit the broad description of a bulbil. Furthermore,
transmission electron microscopic examination of these bulbil
cells reveals that they are engorged with cytoplasmic
structures including lipid bodies and each has a nucleus
(Figs. 5,6)(8,9). They also possess a bilayered cell wall
74
(Figs. 5,6). Nevertheless, these bulbils differ from the
bulbils of other standard bulbilliferous fungi by lacking
pigmentation, certain sclerotial-like qualities (outer rind-
like cells and inner swollen cells) and in the manner of
their formation (10).
T. andreanae may be related to Cladorrhinum spp. by virtue of
certain conidial and hyphal characteristics (11). However,
our SEM and culture observations made on the type species -
Cladorrhinum foecundissimum revealed the absence of bulbils, the
presence of spherical conidia, and the presence of small,
numerous phialid-like structures on the hyphae, presumably
from which the spores are budded (11). Furthermore, C.
foecundissimum and related species are heavily pigmented (green-
black colonies) and are commonly isolated from soil and dung
sources. In addition, T. andreanae was inhibitory to C.
foecundissimum. These findings are all in sharp contrast to the
descriptions given in this report of T. andreanae.
Type species: Taxomyces andreanae Strobel, Stierle, & Hess.
Holotypus: Based on material taken from the bark of Taxus
brevifolia Nutt., collected by Andrea and Don Stierle, August,
1991, in Flathead County, Montana, USA. Agar slants
containing the type culture are deposited with the MSU
mycological collection, D. E. Mathre, Department of Plant
Pathology, Montana State University - col. no. 738.
Duplicate cultures are deposited at the CBS, Baarn,
Netherlands; International Depository Authority, CBS 279.92.
Dried specimens are on deposit at the Farlow Herbarium,
Harvard University (FH).
Etymology
The source tree of the fungus is Taxus, hence the genus
designation - Taxomyces. The species name, Taxomyces andreanae,
is in honor of Andrea Stierle. Andrea is a colleague,
research associate and dedicated scientist. She and her
husband, Don Stierle, were the people who originally isolated
this organism.
Cultural Characterization
When an agar plug of inoculum was placed in the center
of most freshly prepared agar plates enriched with various
nutrients it grew so rapidly that it reached the edge of the
plate in 3 days (corn meal agar, lima bean agar, nutrient
agar, malt agar, oatmeal agar). Bulbils did not form on any
of these media up to 6 days after inoculation. However
bulbils were noticed on the inoculum piece on the corn meal
agar after 6 days. Some bulbils were noticed at the edges of
ihe)
the malt agar plate 7 days after inoculation. Numerous
fluffy aerial mycelia were especially observed on malt agar
and after 6-7 days the mycelium on the malt agar developed a
deep reddish-brown coloration and a thick mycelial mat.
When the fungus was placed on the autoclaved leaves,
fragments of small limbs, and bark taken from various tree
species located in the geographical area of Taxus brevifolia, the
best mycelial growth and bulbil formation occurred on Pacific
yew (Table 1, Fig. 4), followed by River birch (Betula nigra)
(Table 1). In contrast, there was no growth on, or bulbil
formation on, Larix occidentalis, or Tsuga heterophylla, (Table 1).
Other species differentially supported weak fungal growth and
light bulbil formation, eg. Pinus monticola, Picea engelmanni (Table
1). These observations suggest the likelihood that some host
preference of Taxomyces andreanae exists in nature and that it
would be unlikely to be found in and on many species other
than Taxus or Betula. This organism appears to be a saprophyte
or endophyte with the latter preferred since it was found in
association with living tissue. After one year it was still
possible to recover the organism from the inner bark of the
source tree. There is no evident gross pathology of this
host tree. Furthermore, attempts, using agar blocks infested
with T. andreanae placed under the bark of yew also failed to
cause any disease manifestation.
We also noted that the thicker hyphae ca. 3.75 um in
dia, were the ones that typically extended the mycelial mat
from one object (leaf or limb fragment or agar block) to
another. These might be considered "exploratory hyphae".
Careful study of the cultural, mycelial and bulbil
characteristics in comparison to other bulbilliferous fungi
nicely demonstrated the uniqueness of Taxomyces andreanae (10).
ACKNOWLEDGEMENTS
We acknowledge the financial assistance of the National
Science Foundation and the Montana Agricultural Experiment
Station. Appreciation is also expressed to the Hungry Horse
Ranger District of the Flathead National Forest and the
Superintendent of Glacier National Park. We also acknowledge
discussions with Dr. Amy Rossman, ARS, USDA; Dr. Don Mathre
and Gene Ford of Montana State Univesity; and Dr. Ed Butler,
University of California, Davis. The description was
rewritten in taxonomic format and translated by S. L. Welsh
of Brigham Young University. The authors appreciate the help
of Ms. Pam Berger in typing and laying out this manuscript.
Fig. 1. A scanning electron micrograph of hyphae and
fructigenous hyphae of T: andreanae. Bar=10 um.
Fig. 2. A scanning electron micrograph of a series of various
shaped bulbils of T. andreanae. Bar=10 um.
Fig. 3. A scanning electron micrograph of a single bulbil of
T. andreanae illustrating the organization of the cells of the
bulbil. Bar=10 pm.
Fig. 4. Growth of T. andreanae on the inner bark of Pacific yew
(left) and growth and bulbil formation on the leaves and
small limb fragments of T. andreanae (right).
78
Transmission electron micrograph of bulbil cells
of T. andreanae illustrating a dense cytoplasm in
each cell. Bar=1 pm.
Transmission electron micrograph of TI. andreanae
bulbil cells at a higher magnification. Note that
the cell wall has two distinct layers, and the
fiberous material between the cells. Bar=1 um.
79
Table 1. Growth and bulbil formation of Taxomyces andreanae on
various tree species normally growing in the
vicinity of Taxus brevifolia. All observations were
recorded 1 week after inoculation with a 1.0x1.0
agar block supporting fungal growth.
Plant Species Av growth from Formation
edge of agar of
block (cm) bulbils*
Twigs/leaves Bark Twigs/leaves Bark
Taxus brevifolia Nutt.
Pacific Yew 2.0 0.25-0.5 heavy heavy
Betula nigra L.
River Birch 1.0 0.75 heavy moderate
Pinus monticola Doug].
Western white pine 0.1 0.2 light light
Tsuga heterophylla
Rafn Sarg. 0.0 O71 none none
Western hemlock
Pseudotsuga taxifolia
(Poinjebritt: Ou 0 none none
Douglas fir
Thuja plicata Donn.
Western Red Cedar 0125 075 light to heavy
moderate
Picea engelmanni
Parry exEngelm. 0.1 0 moderate none
Engelmann spruce
Larix occidentalis
Nutt. 0 0 none none
Western larch
*Bulbil formation is given in terms of heavy (completely
covering the area where mycelium is growing) to moderate, to
light (little or sparce bulbil formation).
80
REFERENCES
1) WANI, M. C., H. L. TAYLOR, M. E. WALL, P. COGGON, and A.
T. McPHAIL. LO Fass Plant antitumor agents, VI. The
isolation and structure of taxol, a novel antileukemic
and antitumor agent from Taxus brevifolia. J. Am. Chem.
OS tego eoteled «
2) ROWINSKY, E. K., L. A. CAZANAVE, and F. C. DONEHOWER.
1990. Taxol: a novel investigational antimicrotuble
agent. J. Natl. Cancersitnst. $2355124/—-1259-
3) UPADHYAY, R., D. KENFIELD, G. A. STROBEL, and W. M. HESS.
1991. Ascochyta cypericola causing leaf blight of purple
nutsedge (Cyperus rotundus). Can. J. Bot. 69:797-802.
4) HESS, W. M. 1966. Fixation and staining of fungus hyphae
and host plant root tissues for electron microscopy.
Stain Technol. 41:27-35.
5) REYNOLDS, E. S. 1963. The use of lead citrate at high pH
as an electron opaque stain in electron microscopy. J.
Cell Biol’) 172208=212..
6) SPURR, A. R. 1969. A low viscosity epoxy resin embedding
medium for electron microscopy. J. Ultrastruct. Res.
20°31-43%
7) DeBARY, A. 1887. Comparative morphology and biology of
the Fungi; mycetozoa and bacteria. (English
translation). Clarendon Press, Oxford.
8) WEBER, D. J. and W. M. HESS. 1975). Diverse Spores of
Fungi IN Spores Vol VI. Am. Soc. of Microbiol. p. 97-
111.
9) HESS, (Wo oM: 1981. Fungal Organelles and Other Cell
Structures IN The Fungal Spore Morphogenetic Controls,
G. Turian and H. R. Hohl (eds) Academic Press, N.Y. pp
21-41;
10) WERESUB, L. K., and P. M. LeCLAIR. 1971. On Papulaspora
and bulbilliferous basidiomycetes Burgoa and Minimedusa.
Cans. Bot 4932 208-221 3:.
11) COMPENDIUM OF SOIL FUNGI. 1980. Cladorrhinum (Sacc and
Marchal), [{K. H. Domsch, W. Gams and T-H Anderson eds. ]
Academic Press, N.Y.
MY COTAXON
Volume XLVI, pp. 81-85 April-June 1993
TWO NEW SOUTH AMERICAN SPECIES OF
CORTICIACEAE (BASIDIOMYCETES)
WITH AMYLOID SPORES
KURT HJORTSTAM
Malaregatan 12
S-441 35 Alingsas, Sweden
and
LEIF RYVARDEN
University of Oslo, Department of Biology
Division of Botany
Box 1045 Blindern, N-0316 Oslo, Norway
Abstract
Two new South American species of Corticiaceae (Basidiomycetes), are described:
Amyloathelia aspera Hjortst. & Ryv., with asperulate spores from Chile and
Aleurodiscus exasperatus Hjortst. & Ryv. from Brazil.
Amyloathelia aspera Hjortst.& Ryv. sp.nov. Fig. 1.
Basidioma resupinatum, effusum, laxe adnatum, tenue, pelliculare.
Hymenophorum leve, album vel cremeum, postea lutescens; margine indeterminato.
Systema hyphale monomiticum; hyphae basales hyalinae, tenuitunicatae vel
plerumque crassitunicatae, plus minus rectae, leves, 5-6 pm latae; hyphae
subhymeniales tenuitunicatae, circiter 4 pm latae; hyphae omnes fibulatae.
Cystidia nulla. Basidia subclavata vel plerumque tubularia,
tenuitunicata, constricta et sinuosa, 30-35(-40) x 5-7 pm, 4 sterigmatibus. Sporae
subglobosae, crassitunicatae, asperae,
(6-)7-8(-9.5) x (5.5-)6.-6.5(-7) um, amyloideae, cyanophilae.
Holotypus: Chile, Osorno, P.N. Puyehue, Antillanca, 1500 m., on dead branch in
Nothofagus and Drimys winteri forest, 15 April 1988, Rajchenberg No. 4185
(BAFC). Isotypi: GB and O.
BASIDIOME resupinate, effuse, loosely adnate, rather thin and pellicular.
HYMENOPHORE smooth, at first white to cream-coloured, then pale yellowish,
margin indeterminate or usually paler than the fertile part.
HYPHAL SYSTEM monomitic; basal hyphae thin-walled to more commonly with a
slight wall thickening or even thick-walled next to the substratum, hyaline, rather
straight and long-celled, smooth, often branching at right angles, approximately 5-6
um wide, subhymenial hyphae comparatively short-celled, thin-walled, about 4 pm
wide, all hyphae with clamp-connections.
82
Fig. 1. Amyloathelia aspera A) part of the hymenium, B) hyphae from the subiculum,
C) spores, D) SEM of the spores, bar = 1 um. From the holotype.
83
CYSTIDIA absent.
BASIDIA subclavate or when fully developed almost tubular, thin-walled throughout,
constricted and sinuous, 30-35(-40) x 5-7 pm
with four rather long (6-8 pm) sterigmata and a basal clamp-connection.
SPORES subglobose, thick-walled, often with oil drops in the protoplasm, appearing
completely smooth in Cotton blue, but slightly rugose in KOH as well as in Melzer’s
reagent, (6-)7-8(-9.5) x (5.5-)6-6.5(-7) pm, distinctly amyloid and cyanophilous.
The species is easily recognized by its thick-walled, asperulate spores. Its closest
relative seems to be Amyloathelia amylacea (Bourd.& Galz.) Hjortst. which,
however, has smooth, slightly longer spores and with a stronger amyloidity. The
basidia of the latter are in general also shorter.
It should be noted that Corticium vallum G.H. Cunn. was stated to be the same as A.
amylacea by Staplers (1985). This must be a misinterpretation of A. amylacea since
Stalpers described Cunningham’s species with gloeocystidia. A. amylacea (Corticium
amylaceum) has no cystidia, only occasional hyphal ends between the basidia.
Original material of Corticium vallum has not been studied in this case.
Aleurodiscus exasperatus Hjortst. & Ryv. nov.sp. Big
Basidioma resupinatum, arcte adnatum, tenue, quasi 0.1-0.2 mm crassum, plus minus
coriaceum. Hymenophorum leve vel leviter rugosum, primum cremeum tum griseum
vel dilute brunneolum. Systema hyphale monomiticum; hyphae distinctae,
tenuitunicatae, modice ramosae, (2.5-)3-3.5 pm latae, sine fibulis. Gloeocystidia
generaliter 50-80 x 8-10(-12) pm, tenuitunicata, sulphopositivan contentia.
Dendrohyphidiae numerosae, inclusae vel plerumque projectae, ramosissimae, leviter
incrustatae. Basidia clavata, grandia, generaliter 70-100 x 12-15 pm, 4 sterigmatibus.
Sporae ellipsoideae vel aliquantum fusiformes, tenuitunicatae vel leviter
crassiusculae, (15-)20-23 x 8-10 pm, exasperatae, amyloideae.
Holotypus: Brazil, Sao Paulo, Santos, Cananeia, Ilha do Cardoso, 2-5 Feb. 1987,
Hjortstam 16760. (K).
Other specimen examined: Argentine: Tucuman, Ruta 38, km 19, no date indicated,
leg. H. Gomez. Herb. BAFC 24547.
BASIDIOME resupinate, closely attached to the substratum and hardly detachable,
thin to moderately thick, approximately 0.1-0.2 mm thick, more or less coriaceous,
confluent.
HYMENOPHORE smooth or slightly pulverulent, cream-coloured or with a greyish
tint, with age pale brown. Margin indeterminate.
HYPHAL SYSTEM monomitic; hyphae thin-walled, moderately branched, (2.5-)3-3.5
um wide, forming a rather dense subiculum, all hyphae without clamp-connections.
GLOEOCYSTIDIA generally numerous, with granular content, 50-80 x 8-10(-12) p",
blackish in sulphovanillin, thin-walled, with an obtuse tip, basally often with several
protuberances.
DENDROHYPHIDIA numerous, projecting slightly above the basidia, richly
branched, encrusted.
BASIDIA clavate, large, 70-100 pm long or sometimes longer, with a width of 12-15
um just below the four stout sterigmata, without a basal clamp-connection.
Fig. 2. Aleurodiscus exasperatus A) part of the hymenium, B) dendrohyphidia, C)
acanthophyses, D) spores. From the holotype.
85
SPORES ellipsoid to somewhat fusoid, thin-walled or with slight wall thickening,
(15-)20-23 x 9-12 pm, appearing smooth in KOH but slightly rugose in Melzer’s
reagent, amyloid.
Aleurodiscus exasperatus is similar to A. aurantius (Fr.) Schroet., but differs
primarily by its spore shape and gloeocystidia. In the latter species the spores are
ellipsoid, never fusoid and distinctly echinulate to verrucose. Further more, the
gloeocystidia are moniliform at the apex. Macroscopically, the new species is
reminiscent of species of Vuilleminia Maire, but easily separated from species of this
genus by the amyloid spores and sulfo-positive gloeocystidia.
Acknowledgements.
We acknowledge assistance from Dr. R. Watling, Royal Botanic Gardens, Edinburgh,
Scotland who has read the manuscript and suggested improvements.
References.
Stalpers, J.A. 1985. Type studies of the species of Corticium
described by G.H. Cunningham. N.Z.Journ.Bot.
23:301-310.
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MYCOTAXON
Volume XLVII, pp. 87-92 April-June 1993
DEMATIACEOUS HYPHOMYCETES ON FREYCINETIA
(PANDANACEAE). 4. CRYPTOPHIALE
E.H.C. McKENZIE
Herbarium PDD, Landcare Research
Private Bag 92170, Auckland, New Zealand
and
A.J. KUTHUBUTHEEN
Department of Botany, University of Malaya,
59100 Kuala Lumpur, Malaysia
Abstract
Two new species of Cryptophiale are described and figured, C. caudata from
Malaysia and C. novae-caledoniae from New Caledonia. These fungi were
found during a study of dematiaceous hyphomycetes on dead leaves of Fr-
eycinetia.
Introduction
During an examination of over 70 samples of dead Freycinetia leaves from
nine countries (McKenzie 1991), only two collections of Cryptophiale were
found. Both of these, one from New Caledonia and the other from Malaysia,
are distinct new species and are described and figured in this paper.
Cryptophiale caudata McKenzie et Kuthubutheen sp. nov. Fig. 1
Coloniae conspicuae, effusae. Conidiophora e cellulis atro-fuscis, macrone-
matosa, mononematosa, singula, eramosa, erecta, apicem versus parum
curvata, laevia, septata, brunnea, apice acuta, usque ad 230 »m longa, ad
basim 7-10 ym lata, ad apicem 5-7.5 ym. Zona fertilis subapicalis, scutel-
liformis, 45-60 »m longa, 15-24 .m crassa; cellulae conidiogenosae scutello
cellarum sterilium pallide fusco celatae; cellulae steriles 2.5-5 .m latae,
extendentes 6-9 um ultra conidiophororum parietem. Conidia hyalina, ad-
herentia, clavata vel fusiformia, subulata, laevia, guttulata, eseptata, (20-)
23-26 (-28) x 4-4.25 zm, apicem versus 10-25 ym longa, basi obconice trun-
cata.
Figure 1 Cryptophiale caudata, conidiophores and conidia (PDD 60584 -
type). Specimen mounted in hydrous lactophenol.
89
In foliis mortuis pandanaceae Freycinetia sp.
Holotypus PDD 60584.
Colonies conspicuous, effuse. Conidiophores arising from dark brown, smo-
oth, swollen cells, macronematous, mononematous, single, unbranched,
erect, slightly curved towards the apex, smooth, septate, brown, abruptly
tapered to an acute apex, up to 230 »m long x 7-10 «mm thick near the base,
tapering to 5-7.5 um near the apex. Fertile region subapical, 45-60 »m
long, 15-24 »m wide. Conidiogenous cells obscured by a shield of sterile,
pale brown cells; each shield cell 2.5-5 um wide, extending 6-9 um beyond
the conidiophore wall. Conidia produced in slime and adhering to the fertile
part of the conidiophore in a lateral droplet; hyaline, clavate or fusiform,
subulate, smooth, guttulate, not appearing septate, (20-) 23-26 (-28) x 4-
4.25 «.m, tapering to form a single, 10-25 ym long flexuous appendage at
the distal end, base long, obconically truncate.
On Freycinetia sp.
Specimen examined: Malaysia, Selangor, Genting Highlands, Gunung
Bunga Buah, 18.VIII.1992, E.H.C. McKenzie (PDD 60584 - holotype).
The specific epithet refers to the tail-like appendage on the conidia.
Cryptophiale novae-caledoniae McKenzie et Kuthubutheen sp. nov.
Bigio2
Coloniae conspicuae, effusae. Conidiophora e cellulis atro-fuscis, macrone-
matosa, mononematosa, singula, eramosa, erecta, recta vel flexuosa, lae-
via, septata, obscure rubro-brunnea, apicem versus parce pallidiora, acuta,
usque ad 450 ym longa, ad basim 10-11 ,m lata, ad apicem 4-6.5 ».m. Zona
fertilis apicalis, scutelliformis, 35-80 «.m longa, 13-14 ym crassa; cellulae
conidiogenosae scutello cellarum sterilium pallide fusco celatae; cellulae
steriles 2.5-3.5 «m latae, extendentes 3-4 »m ultra conidiophororum pari-
etem. Conidia hyalina, adherentia, fusiformia, laevia, medio septata, (9.5-)
14-16 (-18) x 2-2.25 um, apice rotundata vel acuta, basi rotundata vel ob-
conice truncata.
In foliis mortuis pandanaceae Freycinetia sp.
Holotypus PDD 60585.
Colonies conspicuous, effuse. Conidiophores arising from dark brown, smo-
oth, swollen cells, macronematous, mononematous, single, unbranched,
erect, straight or flexuous, smooth, septate, dark reddish-brown, slightly
paler towards an acute apex, up to 450 »m long x 10-11 ym thick near
the base, tapering to 4-6.5 «m in the fertile region. Fertile region apical,
35-80 »m long, 13-14 wm wide. Conidiogenous cells obscured by a shield of
sterile, pale brown cells; each shield cell 2.5-3.5 um wide, extending 3-4 ».m
beyond the conidiophore wall. Conidia produced in slime and adhering to
90
BrceeadaeAre ay
oT yuna canis HES
FOB BEDII0+60C2: 0-255 eS ne
pe Re == Soe Se
Se
<= fees es
conidiophores and conidia (PDD
d in hydrous lactophenol.
1ae,
-caledon
Figure 2 Cryptophiale novae
60585 - type). Specimen mounte
91
the fertile part of the conidiophore in a lateral droplet; hyaline, fusiform,
smooth, medianly 1-septate, (9.5-) 14-16 (-18) x 2-2.25 ,m, apex rounded or
acute, base rounded or obconically truncate.
On Freycinetia sp.
Specimen examined: New Caledonia, Mt Panié, 1500m, 15.XII.1990, J.S.
Dugdale (PDD 60585 - holotype).
The specific epithet refers to the country where the species was found.
Sutton et al. (1989) provided a key to nine species of Cryptophiale then
known; a tenth species, C. secunda Kuthubutheen & Sutton having been
previously transferred to Cryptophialoidea Kuthubutheen & Nawawi (19-
87). McKenzie (1993) has recently described a new species, C. insularis,
from New Zealand. C. caudata and C. novae-caledoniae can be distin-
guished from other species in the genus by the structure of the conidiophore
apex, size of conidia, and position of the fertile region on the conidiophore.
C. caudata is similar in several respects to C. aristata Kuthubutheen &
Sutton (1985) and to C. enormis Sutton, Nawawi & Kuthubutheen (1989).
The apex of the simple, setiform conidiophore in all three species extends
not more than 10 »m beyond the fertile region. The conidiophore of C.
enormis, however, is wider at the fertile region than above or below it.
The conidia in all three species are caudate. The conidia of C. aristata are
23-27 x 2-3 «.m and 1-septate and the apical cell is drawn into a filiform
appendage 7-12 »m long. Although the conidial length of C. caudata and
C. aristata is similar, in C. caudata the conidia are broader, 0-septate, and
the apical flexuous appendage is 10-25 »m long. The conidia of C. enormis,
however, are twice as large as those of C. caudata, 4-7 septate, and have
shorter (5-8 .m) appendages.
C. novae-caledoniae is reminiscent of C. kakombensis Pirozynski (1968), a
species widely distributed in the tropics and subtropics. The conidia of C.
kakombensis, however, are falcate, longer but narrower (22-28 x 1-2 ,m),
and acute at both ends compared to the conidia of C. novae-caledoniae. The
fertile region of C. kakombensis is subapical and not apical as in C. novae-
caledoniae. The apical location of the fertile region in C. novae-caledoniae
is unique and not found in any of the known species in the genus.
Acknowledgment
We gratefully acknowledge the assistance of Dr G. Kuschel, Auckland, in
preparation of Latin diagnoses. We thank Professor Gareth Morgan-Jones,
Department of Plant Pathology, College of Agriculture, Auburn University,
Alabama, USA for critical review of this paper.
92
References
Kuthubutheen, A.J.; Nawawi, A. 1987: Cryptophialoidea gen. nov. on
decaying leaves from Malaysia. Transactions of the British Mycological
Society 89: 581-583.
Kuthubutheen, A.J.; Sutton, B.C. 1985: Cryptophiale from Malaysia. Tran-
sactions of the British Mycological Society 84: 303-306.
McKenzie, E.H.C. 1991: Dematiaceous hyphomycetes on Freycinetia (Pan-
danaceae). 1. Stachybotrys. Mycotaxon 41: 179-188.
McKenzie, E.H.C. 1993: New hyphomycete species from litter in the Chat-
ham Islands, New Zealand. Mycotaxon 46: 291-297.
Pirozynski, K.A. 1968: Cryptophiale, a new genus of hyphomycetes. Cana-
dian Journal of Botany 46: 1123-1127.
Sutton, B.C.; Nawawi, A.; Kuthubutheen, A.J. 1989: Additions to Belem-
nospora and Cryptophiale from Malaysia. Mycological Research 92: 354-
358.
MYCOTAXON
Volume XLVII, pp. 93-100 April-June 1993
GENETICS OF A TUBER AESTIVUM POPULATION
(Ascomycotina, Tuberales)
Gi, PACIONE 7 3Ge(ERLZAL 7 M. MIRANDA and C. VISCA
Dipartimento di Scienze Ambientali and i Dip Biologia&
Fisiologia Cellulare, Universita, 67100 L'Aquila, Italy
Key-words: Tuberales, Tuber aestivum, Tuber uncinatum,
truffle, allozymes, reproduction, sibling species.
Abstract: A truffle population with of Tuber aestivum and
Tuber uncinatum morphotypes was subjected to allozyme
analysis via SGE and genotypically scored at fourteen loci.
Ten of the loci were monomorphic and four polymorphic. Thus,
seventeen new genotypic classes, with very low genetic
differentiation, were found. However, heterozygosity was
never found and all alleles seem to be fixed in individuals.
These results indicate that Tuber aestivum and T. uncinatum
are probably two extreme morphological forms of a complex of
self-fertilizing sibling species.
Introduction
Some taxonomic complexes of the genus Tuber Mich. ex Fr.
are very difficult to distinguish on a morphological basis.
For Tuber aestivum Vitt., a European taxon, also found in
Algeria and Turkey, there seemsjito be a’ long list,of
probable synonyms. Beyond the nomenclatural problems, the
difficulty in distinguishing species has produced confusion
in regulating and marketing these species worldwide.
CHEVALIER et al. (1978) in an attempt at’ separating Tuber
uncinatum Chat. as a distinct species, tried without much
success to juxtapose its morphological characters with those
of T. aestivum. GROSS (1987) proposed to divide these two
taxa on the basis of the height of the sporal reticulum.
Moreover, in two previous reports, no heterozygosity was
found for several loci in a number of Tuber species (PACIONI
& POMPONI, 1989; 1991). An investigation at the population
94
level, to test the hypothesis that a selfing process of
reproduction may be responsible for the homozygosity,
therefore appears of some interest.
This paper reports the results of using allozyme analysis to
resolve the status of Tuber aestivum and T. uncinatum.
Materials and Methods
A sample of 66 specimens, morphologically related to Tuber
aestivum and Tuber uncinatum, was collected during 1990 and
1991 ‘inthe, } bei Poz2ze"," aysmail) trutrie preserve; near che
village of Santi, in’ LD Aguilav(The Abruzzi »yltalyysithe
specimens, determined by the senior author, have been
preserved, one portion cut in little sticks and then deep-
frozen and a small portion dried and deposited in the
mycological herbarium of L'Aquila (AQUI) as a voucher.
Horizontal starch gel (SGE) was performed essentially
according to PACIONI & POMPONI (1989) and FRIZZI & TAMMARO
(1991). Some slight modifications were made in the staining
solutions. The stains were basically prepared according to
HARRIS (1966), exceptufor the’ SKDH. stain) (FINESCHI, 1983).
Electrophoresis was conducted at a constant 8V/cm for
approximately 3.5 hours per run.
Details on the electrophoretic techniques used are given in
Wavie Wis
A preliminary survey showed fourteen enzymatic loci
produced patterns clearly interpretable from a genetic point
of view: Pgm-2 (PGM, phosphoglucomutase), G6pd (G6PD,
glucose-6-phosphate dehydrogenase), 6-Pgd (6PGD,
phosphogluconate dehydrogenase), Hk (HK, hexokinase), Mpi-1l
(MPI, mannose phosphate isomerase), Ak (AK, adenylate
kinase), Pgi (PGI, glucose phosphate isomerase), Mdh-2 (MDH,
malate dehydrogenase), Me-1 and Me-3 (ME, malic enzyme),
Idh (IDH, isocitrate dehydrogenase), Sod-1 (SOD, superoxide
dismutase), Skdh (SKDH, shikimic dehydrogenase).
With a view to comparing the resulting elecrophoretic data
to all known data for Tuber, the previous reference for the
mobility of the enzymes of Tuber melanosporum Vitt. (=100)
was mantained.
Electrophoretic data were analysed using the NTSYS-pc
program (ROHLF,1987) and genetic divergence was estimated
with the index of standard distance (D) proposed by NEI
CL9I2)>
Results
Nine of the fourteen enzymatic loci examined ( G6pd,
6Pgd, Adk, Got, Pgi, Me-3, Skdh, Mdh-1, Sod) are monomorphic
95
showing the same phenotype as one band. The Pgm-2 locus is
also monomorphic, however it is distinguished by three
bands. The Pgm-2 pattern has been reported for all other
taxa of Tuber already examined (PACIONI & POMPONI, 1989;
1991). The cause of the Pgm-2 pattern is still unexplained.
The relative mobility of the above enzyme electrophoretic
patterns are: 6Pgd 103, Adk 100, Got 105, Pgi 100, Me-3 110,
Skdh 100, Mdh-1 100, Sod 93, Pgm-2:85. Only LOurgocie(Hk:,
Mpi-1, Me-1, IdH) appeared polymorphic. At locus Hk three
alleles (108; (110; 7112). were found, two. (95),..100:) at. Mpi-1,
two (105, 110), at \Me-l*and’three (90, 95, 100) at.iIdh.
Heterozygosity was not found and all alleles seem to be
fixed individuals.
The electrophoretic patterns revealed seventeen genotypic
classes (ROYSE & MAY, 1982). Only the four polymorphic loci
are reported (Table 2). Their genetic distances are very
LOW ithe SiO), Oi 4 oe OmMEOr 6s SOK) IN LOD Un Ce Vales moti an
intraspecific variability, confirming the overlapping of the
morphocharacters. Figure 1 shows one of the possible
clustering trees. The enzyme phenotype, belonging to the
genotypic class labeled TA2, seemed to be the most common.
Several phenotypes were only observed once. The number of
phenotypes found is almost equal to that of the truffle
habitats sampled.
For loci in common, the genotypic classes found here are
completely different from those previously observed in
samples from other Italian localities (PACIONI & POMPONI,
L991).
Discussion
The homozygosity of truffles has been reported previously
in the studies on the Tuber aestivum-T. mesentericum group
(PACIONI & POMPONI, 1991) as well as on several other taxa
of Tuber (PACIONI & POMPONI, 1989). However vegetative cells
of the truffle sporophores contain a pair of nuclei (FASOLO-
BONFANTE & BRUNEL,1972) and therefore a certain percentage
of heterozygosity should be present, unless the two nuclei
arise from self-fertilization or mitosis.
Our results strongly support the hypothesis that in
dikaryotic hyphae, the two nuclei result from selfing;
accordingly non-heterozygosity is observed. This is at
variance with what is known about truffle reproduction,
hitherto supposed to be heterothallic (DANGEARD 1895/96;
GRENTE et al., 1972).
Sexual reproduction by selfing would restrict, in this group
of) Lungijevariabi laity toh tnter=and intrachromosomal
recombination and mutation. As stated in the Results, only
96
four of fourteen loci examined appeared polymorphic and this
may favour the hypothesis that a strong selection, due to
the, peculiaryjecology; and ‘lute cycles ofs:trufives /ihas> been
operating on the ten monomorphic loci, whereas it has been
less stringent on the polymorphic loci. It should be noted,
however, that, rnytrutites, very little inftormationeis
available about the structures, functions and molecular
properties’ of thetenzymes, anvéestigated here. jin tact, ethe
polymorphism observed might be ascribed to alleles of the
same locus, to alleles of entirely different loci
differentially expressed in the various samples, or to
conformational variants (DAWSON, 1972) of the same locus
expressed in different sporophores.
In the light of the present results, we think that the
sporophores of Tuber aestivum or T. uncinatum represent only
two extreme forms, at present unresolvable by morphological
and biometrical methods.
The existence of a great number of sibling species suggests
that .oun knowledge! otmttber jreproduetionsismstallyvery
scanty and requires more investigation.
Research carried out with financial support from M.U.R.S.T.,
Scientific Research 40% "Ecology and biology of fungi and
lichens.’ We are! andebbedi to Prof. 68.) Grilbis for his
assistance with the English manuscript and Prof. R.D. Fogel,
University of Michigan, for his suggestions.
References
BREWER,G. & SING,C.F., 1970. An introduction to isozyme
techniques. London & New York: Academic Press.
CHEVALIER; -G., DESMAS; C., FROCHOT, H. & RIOUSSET;, L:, 1978.
L'éspece Tuber aestivum Vitt. : I. Definition. Mushroom
SciencelO: 957-975.
DANGEARD, P.A., 1895/96. La Truffle. Recherches sur _ son
développement, sa structure, sa reproduction sexuelle.
Botaniste 4/5:63-87.
FASOLO-BONFANTE,P. & BRUNEL,A., 1972. Caryological features
in a mycorrhizal fungus: Tuber melanosporum. Allionia 18:
5-11.
FPINESCHI) (52% P983kiVariabilitay untraspecificaritiny /einus
nigra, Arn... 4 iRisultatiervd)? Ganalisi ) sugvalcuni ssisteni
isoenzimatici. L'Italia Forestale e Montana 38: 200-213.
FRIZZI, G. & TAMMARO, F.; 1991. Electrophoretic study “and
genetic affinity sin ‘the Campanula elatines and Gx
fragilis (Campanulaceae) rock-plants group from Italy and
W. Jugoslavia. Plant Syst. Evol. 174: 67-73.
o7.
GRENTE,J., CHEVALIER, Geese POLLACSER en 9:7 or La
germination de l'ascospore de Tuber melanosporum et de la
synthése sporale des mycorrhizes. C.R. Acad. Sc. Paris
21 Ot ea 3.
GROSS, G., 1987. Zur den europaischen Sippen der Gattung
Tuber. in,DERBSCH,H.,& SCHMITT, J.A., (Eds.):Atlas der
Pilze des Saarlandes,2: Nacweise, Okologie, Vorkommen und
Beschreibungen, pp.79-99. Saarbrucken, Germany: Ed.
Delattinia.
HARRIS, H., 1966. Enzyme polymorphism in man. Proc. Roy.
soc. London B69 *% 3298-31107
NEI, M., 1972. Genetic distance between populations. Americ.
NaG. 61062 (233-292.
PACLONI7: 9G. &, POMPONI, G:, "1989. ‘Chemotaxonomy of ‘some
Italian species of Tuber. Micol. Veget. Medit. 4: 63-72.
PACIONI, G. & POMPONI, G., 1991. Genotypic patterns of some
Italian populations of the Tuber aestivum-T. mesentericum
complex. Mycotexon 42:3, 171-179.
ROHLF, estllinan 1987. NTSYS-pc Numerical Taxonomy and
Multivariate Analysis System. Setauker, N.Y., U.S.A.:
Exeter Publishing.
ROTHE, G., 1973. Shikimisaure-Dehydrogenase in keimenden
Erbsen. Biochem. Biophys. Pflanzen 164: 475-486.
ROYSE, 7 Di.- & MAY, 9B.) 61982. Use of, tseozyme: ‘variation to
identify genotypic classes of Agaricus brunnescens.
Mycologia 74: 93-102.
SiOUANDER gk. Ws 2 OM LIn. jo MH. @YANG Sy, GONHSON, Wok. o&
GENTRY, J.B., 1971. Biochemical polymorphism in the genus
PeromnyScusi tl. a Variation pin. ene old field mouse
(Peromyscus polionotus). Stud. Genet. 6: 49-90.
SHAW, C.R. & PRASAD, R., 1970. Starch gel electrophoresis of
enzymes. A compilation of recipes. Biochem. Genet. 4:
297-320.
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MY COTAXON
Volume XLVI, pp. 101-129 April-June 1993
NEW SPECIES IN THE LICHEN FAMILY PARMELIACEAE
(ASCOMYCOTINA) FROM AUSTRALIA
JOHN A. ELIX
Department of Chemistry, The Faculties, Australian National University,
GPO Box 4, Canberra, ACT, 2601, Australia
ABSTRACT : The species, Bulbothrix microscopica
Elix, Canoparmelia herveyensis Elix, Canoparmelia
subarida Elix, Canoparmelia whinrayi Elix,
Flavoparmelia kantvilasii Elix, Hypotrachyna
booralensis Elix, Hypotrachyna proserpinensis Elix,
Hypotrachyna subpustulifera Elix, Neofuscelia
kondininensis Elix, Neofuscelia imitatricoides Elix,
Neofuscelia parasitica Elix, Neofuscelia scabrosina
Elix, Neofuscelia subbarbatica Elix, Paraparmelia
yamblaensis Elix, Parmelia crowii Elix, Parmelina
euplectina Elix, Parmelina johnstoniae Elix,
Parmelinopsis protocetrarica Elix, Parmotrema
submerrillii Elix, Xanthoparmelia austroconstrictans
Elix, Xanthoparmelia canobolasensis Elix,
Xanthoparmelia hyposalazinica Elix, Xanthoparmelia
masonii Elix and Xanthoparmelia trirosea Elix are
described as new. A new name is given to
Xanthoparmelia xanthofarinosa Elix (formerly xX.
xanthosorediata). The following new combinations
are made: Bulbothrix queenslandica (Elix & Stevens)
Elix, Bulbothrix subtabacina (Elix) Elix, Canoparmelia
owariensis (Asah.) Elix, Canoparmelia pustulescens
(Kurok.) Elix, Canomaculina melanochaeta (Kurok.)
Elix, Imshaugia evernica (Elix & Johnst.) Elix,
Parmotrema pseudovirens (Gyelnik) Elix and
Rimeliella haitiensis (Hale) Elix.
Introduction
As a result of further study of the lichen family Parmeliaceae in
Australia (Elix & Johnston 1986, 1987a, 1987b, 1987c, 1988a, 1988b, 1988c; Elix,
Johnston & Armstrong 1986; Nash & Elix 1987), I am describing 24 new
species. Throughout the present work chemical constituents were
identified by thin layer chromatography ( Culberson 1972; Culberson,
Culberson & Johnson 1981; Culberson & Johnson 1982; Elix, Johnston &
102
Parker 1987, 1988), high performance liquid chromatography (Lumbsch &
Elix 1985) and comparison with authentic samples.
Bulbothrix microscopica Elix sp. nov. Fig. 1
Species cum thallo ut in Bulbothrix subtabacina sed ab hac specie
thallo diminuto, lobis angustioribus, emaculatis et ciliis inflatis differt.
Type: Australia. Queensland. On mangroves along the foreshore of Tin
Can Inlet; Tin Can Bay, 25°54'S, 153°0V'E, Tom iy. Av Elix 22862) 3.vili 1989;
holo: CBG.
Thallus corticolous, adnate to tightly adnate, mineral grey, 1-2 cm
diam. Lobes sublinear, 0.1-0.5 mm wide, subdichotomously branched,
margins densely bulbate-ciliate, bulbate cilia shiny, markedly inflated.
Upper surface plane, emaculate, moderately to densely isidiate, the isidia .
simple, less than 0.15 mm high. Medulla white. Lower surface black shiny,
moderately to densely rhizinate to the margins, rhizines black, simple or
tufted at apices. Pycnidia and apothecia not seen.
Chemistry. Cortex K+ yellow, medulla K+ yellow-red, C-, P+ orange;
containing atranorin, chloroatranorin, salazinic acid (major), consalazinic
acid (minor).
The minute thallus, very narrow lobes, simple isidia, black lower
surface and medullary salazinic acid distinguish this species. P. subtabacina
Elix a related species (vide infra), has broader (0.5-1.0 mm wide), sublinear-
elongate lobes with a distinctly maculate upper surface, slender, elongated
bulbate cilia which are commonly branched subapically and dichotomously
branched rhizines. Currently this rare endemic species is only known from
the type locality.
Canoparmelia herveyensis Elix sp. nov. Fig. 2
Species cum thallo ut in Canoparmelia raunkiaeri sed ab hac specie
medulla inferiore pigmento ochraceo et euplectino continente differt.
Type: Australia. Queensland. On granite rocks in dry sclerophyll forest,
Hervey Range, 45 km south-west of Townsville, 19°26'S, 146°24’E, 350 m, J.
A. Elix 20427 & H. Streimann, 20.vi.1986; holo: CBG; iso: MEL.
Thallus adnate, saxicolous, pale mineral grey, 5-8 cm diam. Lobes
subirregular, apically rotund, 2-4 mm wide. Upper surface plane, becoming
irregularly rugose towards the centre, pustulate-sorediate, the pustules
coalescing in a coarsely sorediate mass in the older portions of the thallus.
Medulla white for the most part but pigmented yellow-orange to orange-
brown adjacent to lower surface. Lower surface black, sparsely rhizinate,
103
rhizines simple, with a narrow bare, brown zone at the margins. Apothecia
and pycnidia not seen.
Chemistry. Cortex K+ yellow, medulla K-, C-, KC-, P+ orange-red, lower
medulla K+ violet; containing atranorin, chloroatranorin, protocetraric acid
(major), virensic acid (trace), euplectin (minor).
This new species resembles Canoparmelia raunkiaeri (Vainio) Elix &
Hale as both develop pustulate soredia towards the centre of the thallus and
contain medullary protocetraric acid. However, C. herveyensis, is clearly
distinguished by the partially orange-yellow pigmented lower medulla due
to the presence of the anthraquinone, euplectin. At present this new species
is only known from the type locality where it appears quite common.
Canoparmelia subarida Elix sp. nov. Figs 3
Species cum thallo ut in Canoparmelia macrospora sed ab hac specie
sporis brevioribus et acido lecanorico continente differt.
Type: Australia. Western Australia. On twigs, South Stirling sand plain, 4
km off the Chillinup road, N. Sammy UWA8, 12.viii.1970; holo: PERTH.
Thallus corticolous, adnate, pale mineral grey, 1-3 cm in diam. Lobes
subirregular, crowded and imbricate, 1.0-2.0 mm wide. Upper surface
smooth and shiny at the apices, plane or weakly foveolate at first, becoming
wrinkled and rugose with age, lacking soredia and isidia. Medulla white.
Lower surface ivory to pale brown, moderately to densely rhizinate, rhizines
concolorous with the lower surface, simple or tufted at the tips. Pycnidia
numerous, punctiform, black; conidia cylindrical 5-6 x 1 um. Apothecia
common, sessile, 0.5-2 mm in diam., disc concave at first but becoming
flattened with age, chestnut brown to dark brown, epruinose; margin thin,
entire, smooth; spores (8), colourless, ellipsoid, 7-9 x 5-6 um.
Chemistry. Cortex K+ yellow, medulla K-, C+ red, P- ; containing atranorin,
chloroatranorin, lecanoric acid (major), orsellinic acid (trace), unknown
(trace).
This species is one of the few species of Canoparmelia with a pale
lower surface. In gross morphology C. subarida closely resembles C.
macrospora Elix & Johnston, but the latter species is distinguished by the
maculate apical lobes which are densely pruinose, the large (16-20 x 5-6 um)
often curved spores and medullary scabrosin 4,4'-diacetate. C. subarida is
chemically identical to C. pruinata, but the latter species is distinguished by
the much larger (4-15 mm), pruinose apothecia and a black lower surface. C.
subarida appears to be a rare species in the subarid inland areas of southern
Australia.
104
Figures 1-4. New species of Parmeliaceae: 1, Bulbothrix microscopica
(holotype in CBG); 2, Canoparmelia herveyensis (holotype in CBG); 3,
Canoparmelia subarida (holotype in PERTH); 4, Canoparmelia whinrayi
(holotype in MEL). Scale bar = 5 mm.
105
Specimens Examined
VICTORIA. On twigs, W side of Wyperfeld National Park, along fire access
trail, c. 31.55 km W of Wonga Hut, R. Filson 16024, 29.viii.1978 (MEL).
SOUTH AUSTRALIA. On bark of Acacia aneura, 20 km W of Vokes
Corner, 28°34'S, 130°29'E, N. N. Donner 7376, 23.viii.1980 (AD).
Canoparmelia whinrayii Elix sp. nov. Fig. 4
Species cum thallo ut in Parmelina labrosa sed ab hac specie thallo
subto pallido et cilia desunt differt.
Type: Australia. Tasmania. On twigs, north-west of Trig Point, Flinders
Island, John Whinray, 1.i1.1969; holo: MEL.
Thallus corticolous, adnate, olive-grey to mineral grey, c. 7 cm diam.
Lobes subirregular and apically subrotund, 3-5 mm wide, lacking cilia.
Upper surface plane to wrinkled, foveolate, shining or rather waxy,
sorediate; soralia laminal or marginal, white, becoming grey-brown with
age, originating from coarse pustular ridges, spreading over the thallus.
Medulla white. Lower surface ivory to pale tan, moderately rhizinate,
rhizines simple, concolorous with lower surface. Pycnidia and apothecia
not seen.
Chemistry. Cortex K+ yellow, medulla K-, C+ red, P-; containing atranorin,
chloroatranorin, lecanoric acid (major).
This unique new species is distinguished by the broad lobes with a
pale lower surface, the pustular soralia, the medullary lecanoric acid and the
absence of pseudocyphellae and cilia. It is the only sorediate Canoparmelia
with a pale lower surface so far recorded for Australia. In overall growth
form and in the development of soralia, this new species resembles
Parmelina labrosa (Zahlbr.) Elix & Johnston, but the latter is distinguished
by the sparse marginal cilia and the black lower surface. At present this
species is only known from the type collection.
Flavoparmelia kantvilasii Elix sp. nov. Fig. 5
Species cum thallo ut in Flavoparmelia euplecta sed ab hac specie
thallo diminuto, medulla albida, inferiore nonpigmentifera et euplectino
noncontinente differt.
Type: Australia. New South Wales. On twigs of Lomatia arborescens in
Eucalyptus pauciflora woodland, Thunderbolt Lookout, Barrington Tops
National Park, 25°54'S, 153°01'E, 1450 m, G. Kantvilas 426/88, 4.vii.1988;
holo: HO, iso: NSW, ANUC.
106
Thallus corticolous, adnate, green to yellow-green, 2-4 cm in diam.
Lobes irregularly rotund, incised, margins crenulate, 2.0-5.0 mm wide;
secondary lobes narrower, overlying the marginal lobes. Upper surface
plane, matt, smooth at the marginal lobes becoming slightly rugulose
towards the centre of the thallus, isidia absent, pustulate-sorediose; soralia
sometimes developing from cracks or pustules which burst to form
granular soredia. Medulla white. Lower surface jet black, with a pale brown
bare zone at the lobe ends, sparsely rhizinate; rhizines simple or fasciculate.
Pycnidia and apothecia not seen.
Chemistry. Cortex K-, medulla K-, C-, P+ red; containing usnic acid,
protocetraric acid (major), atranorin (+), caperatic acid, virensic acid (¢ trace),
unknown (+ trace).
Morphologically this species closely resembles Flavoparmelia
euplecta (Stirton) Hale as both species have adnate thalli with a pustulate
sorediate upper surface and contain protocetraric acid as a major medullary
constituent. However the thalli of F. kantvilasii are smaller (2-4 cm cf. 4-12
cm) and more fragmented, not only when growing on its normal twig
habitat but even when found on broader branches. Furthermore F.
kantvilasii lacks the orange pigmented lower medulla (K+ purple,
containing euplectin) so characteristic of F. euplecta. F. kantvilasii is found
in the temperate forests of northern New South Wales.
Specimens Examined
NEW SOUTH WALES. On Ficus in coastal Eucalyptus forest, Telegraph
Point, 25 km NNW of Port Macquarie, 31°19'S, 152°48'E, K. L. Gaul, viii.1986
(ANUC); on canopy twigs of Nothofagus moorei in rainforest, Burraga
Swamp, Mt Allyn Forest Park, Chichester State Forest, 32°06'30"S,
151°25'30"E, 1000 m, G. Kantvilas 187/88, 29.vi.1988 (ANUC, HO, NSW); on
canopy twigs of Nothofagus moorei in rainforest, Barrington Tops National
Park, Mt William, 1400 m, G. Kantvilas 307/88, 30.vi.1988 (HO, NSW); on
canopy twigs of Nothofagus moorei in rainforest, Gloucester Tops, 1150 m,
G. Kantvilas 414/88A; 2.vii.1988 (HO, NSW); on twigs of Nothofagus
moorei in rainforest, Tweed Range, Wiangerie Forest Drive, Antarctic
Beech Walk, 1000 m, G. Kantvilas 703/88, 3.viii.1988 (HO, NSW); on canopy
twigs of Nothofagus moorei, New England National Park, Cascade Creek
near Wright's Lookout, G. Kantvilas 705/88, 6.viii.1988 (HO, NSW).
Hypotrachyna booralensis Elix sp. nov. Fig. 6
Species cum thallo ut in Hypotrachyna densirhizinata sed ab hac
specie thallo multo parviore et lobi angustiore differt.
Type: Australia. New South Wales. On bark, in a bushland gully, 3 km west
of Booral, Rex Filson 16764, 1.ii.1969; holo: MEL.
107
Thallus corticolous, adnate, whitish-grey to mineral grey, c. 2.5 cm
diam. Lobes sublinear, 0.5-0.8 mm wide, dichotomously branched, separate,
the margins subcrenate. Upper surface shiny on younger lobes, dull and
irregularly cracked on older lobes, plane, emaculate, sorediate; soralia
orbicular, mostly subterminal, capitate. Medulla white. Lower surface
black, densely rhizinate, rhizines black, moderately dichotomously
branched. Pycnidia and apothecia not seen.
Chemistry. Cortex K+ yellow, medulla K-, C-, KC+ rose, P-; containing
atranorin, chloroatranorin, alectoronic acid (major).
This new species resembles the American-African species
Hypotrachyna densirhizinata (Kurok.) Hale in that both are sorediate with
capitate soralia and contain alectoronic acid. However H. densirhizinata is
clearly distinguished by the size of the thallus (6-20 cm diam.), the breadth of
the lobes (2-6 mm wide) and the presence of additional a-collatolic acid. At
present this new species is only known from the type collection.
Hypotrachyna proserpinensis Elix sp. nov. Fig. 7
Species cum thallo ut in Hypotrachyna heterochroa sed ab hac specie
thallo laxe adnato, medulla albida, inferiore nonpigmentifera et acido
gyrophorico continente differt.
Type: Australia. Queensland. On granite rocks on edge of escarpment in
Eucalyptus-Casuarina dominated woodland, Clarke Range, 46 km south of
Proserpine, 20°50'S, 148°32'E, 800 m, J. A. Elix 20929 & H. Streimann,
29.vi.1986; holo: CBG; iso: MEL.
Thallus saxicolous, loosely adnate, whitish mineral grey, 5-9 cm
diam. Lobes sublinear, contiguous, 1-2 mm wide, subdichotomously
branched, apices incised, margins entire, axils sinuous. Upper surface plane,
continuous, soredia and isidia lacking. Medulla white. Lower surface black,
moderately to densely rhizinate, rhizines black, densely dichotomously
branched. Pycnidia scattered, conidia bifusiform 5-7 x 1 um. Apothecia
common, substipitate, 2-4 mm diam., thalline exciple smooth to undulate;
spores (8), colourless, ellipsoid, 8-11 x 5-7 um.
Chemistry: Cortex K+ yellow; medulla K-, C+ pink, P-; containing atranorin,
chloroatranorin, gyrophoric acid (major), 3-methoxy-2,4-di-O-
methygyrophoric acid (trace), 2,4-di-O-methygyrophoric acid (trace),
unknown (minor).
Morphologically this new species resembles Hypotrachyna
heterochroa (Hale & Kurok.) Elix, as both have narrow sublinear lobes
which are commonly fertile and lack soredia and isidia. However whereas
the thalli of H. heterochroa are adnate, have a pigmented lower medulla
108
and contain protocetraric acid and skyrin, H. proserpinensis has loosely
adnate thalli with a white medulla containing gyrophoric acid. At present
this new species is known only from the type collection.
Hypotrachyna subpustulifera Elix sp. nov. Fig. 8
Species cum thallo ut in Hypotrachyna pustulifera sed ab hac specie
thallo saxicolo, multo parviore et lobis angustioribus differt.
Type: Australia. New South Wales. On exposed sandstone rocks in dry
sclerophyll forest, trail to Pigeon House Mountain, 19 km west of Ulladulla,
35°20'S, 150°16'E, 460 m, J. A. Elix 21305 & H. Streimann, 2.xii.1986; holo:
CBG; iso HO, MEL.
Thallus corticolous, adnate to tightly adnate, whitish mineral-grey,
2.5-4.0 cm diam. Lobes linear-elongate, separate to subimbricate,
dichotomous to subirregularly divided, 0.5-1.0 mm wide, margins entire.
Upper surface shiny, continuous, plane to slightly convex, emaculate,
pustulate; pustules subapical, bursting open, eroding and becoming coarsely
granular-sorediate with age and exposing black lower cortex. Medulla
white. Lower surface black, moderately to densely rhizinate below and at
the margins, rhizines black, shiny, dichotomously branched. Pycnidia and
apothecia not seen.
Chemistry. Cortex K-, medulla K- or K+ pale reddish, C- KC+ rose, P-;
containing atranorin, chloroatranorin, colensoic acid (major), 4-O-
methylphysodic acid (major), lividic acid (trace), physodic acid (minor),
oxyphysodic acid (trace).
Superficially this new species resembles Hypotrachyna pustulifera
(Hale) Hale as both species have erumpent pustules on the upper surface
and contain components of the lividic acid-physodic acid chemosyndrome.
However H. pustulifera is corticolous and has much larger thalli (4-6 cm
diam.), broader lobes (2-3 mm wide) and contains lividic acid as the major
secondary metabolite. H. subpustulifera by contrast is saxicolous, has
smaller thalli (2.5-4.0 cm diam.), narrower lobes (0.5-1.0 mm wide) and
contains 4-O-methylphysodic acid and colensoic acid as major secondary
substances. This new species is known from several localities in New South
Wales, growing on Hawksbury sandstone.
Specimens Examined
NEW SOUTH WALES. On sandstone rocks in heath, Peckmans Plateau,
Katoomba, 980 m, J. A. Elix 3211, 24.iv.1977 (ANUC); on sandstone rocks in
open Eucalyptus woodland, Morton National Park, 8 km north-east of
Nerriga, 35°07'S, 150°08'E, 750 m, J. A. Elix 11598 & P. W. James, 20.i.1984
(ANUC).
109
Figures 5-8. New species of Parmeliaceae: 5, Flavoparmelia kantvilasii
(isotype in ANUC); 6, Hypotrachyna booralensis (holotype in MEL); 7,
Hypotrachyna_ proserpinensis (holotype in CBG); 4, Hypotrachyna
subpustulifera (holotype in CBG). Scale bar (Fig. 5,7) = 10 mm; (Fig. 6,8) = 5
mm.
110
Neofuscelia kondininensis Elix sp. nov. Fig. 9
Species cum thallo ut in Neofuscelia incantata sed ab hac specie subto
pallido differt.
Type: Australia. Western Australia. On exposed rocks in open scrubland,
with scattered Acacia and Eucalyptus, 29 km east of Kondinin on road to
Hyden, J. A. Elix 21680 & M. V. Sargent, 19. viii.1987; holo-CBG.
Thallus saxicolous, tightly adnate, yellowish brown to dark brown, 4-5
cm in diam. Lobes sublinear-elongate to subirregular, barely imbricate, 0.5-
1.5 mm wide. Upper surface dull or slightly shining, smooth at the
margins, becoming rugulose on older lobes, isidiate; isidia cylindrical,
simple for the most part, seldom branched. Lower surface ivory to pale
brown, moderately rhizinate, rhizines simple, to 0.3 mm long. Pycnidia
rare, conidia not seen. Apothecia not seen.
Chemistry. Cortex K-, HNO3+ dark blue-green; medulla K-, C-, KC+ rose-
red, P-; containing physodic acid.
This new species closely resembles Neofuscelia incantata (Essl.) Essl.
both morphologically and chemically, but differs in the colour of the lower
surface. In N. incantata the underside is dark brown or black, but often paler
at the apices whereas in N. kondininensis the lower surface is ivory to pale
brown, but becoming darker at the apices. At present this new species is
known only from the type collection.
Neofuscelia imatatricoides Elix sp. nov. Fig. 10
Species cum thallo ut in Neofuscelia imitatrix simili sed ab hac specie
lobis accessoriis saepe subcylindrico et acido 4-O-methylphysodico, acido a-
collatolico et acido alectoronico continente differt.
Type: Australia. New South Wales. On sandstone rocks in open Eucalyptus
forest, Morton National Park, 8 km north-east of Nerriga, 35 07'S, 150 08'E,
760 m, J. A. Elix-3062, 30.iii.1977; CBG-holotype, MEL-isotype.
Thallus foliose, saxicolous, moderately adnate, dark brown to black-
brown, paler at the periphery, 3-4 cm diam. Lobes sublinear-elongate, 0.6-1.0
(-1.2) mm wide, flat or slightly convex at the apices, becoming more strongly
convex, cracked and areolate within the thallus, developing very narrow
(0.1-0.4 mm wide), elongate, dichotomously divided secondary lobes at the
periphery and within the thallus. The secondary lobes subfruticose to
fruticose in habit, circular to subcircular in cross section and for the most
part unattached to the substrate. Upper surface dull to slightly shiny at the
apices, smooth at the apices but becoming markedly rugose within the
thallus, lacking soredia and isidia. Medulla white. Lower surface jet black,
111
shining, sparsely rhizinate, rhizines black, short, simple or occasionaly
tufted, slender. Pycnidia common, conidia bifusiform, 6-8 x 1 um.
Apothecia common, short stipitate, 1-3 mm in diam., disc black-brown,
concave at first but becoming more or less flat or undulating, margin thin,
entire; spores (8), colourless, ellipsoid, 7-9 x 3.5-5.5 um.
Chemistry. Cortex K-, HNO+ dark blue-green, medulla K-, C-, P-; containing
alectoronic acid (major), a-collatolic acid (major), 4-O-methylphysodic acid
(major) and physodic acid (minor).
This new species appears to be related to Neofuscelia imitatrix
(Taylor) Essl. since both species often develop distinctly pulvinate thalli
with a black lower surface, and give similar chemical reactions. However
they can be separated clearly by their morphology since the secondary lobes
of N. imitatricoides are distinctly circular to subcircular in cross section
while those of N. imitatrix are flat to weakly convex. The chemical profile
of these two species also differs consistently since N. imitatricoides contains
alectoronic acid, a-collatolic acid and 4-O-methylphysodic acid (all in
substantial proportions) and physodic acid (traces or minor) while N.
imitatrix contains physodic acid (major) together with occasional traces of 4-
O-methylphysodic acid. Thus these two species appear to have overlapping
chemistries (i.e. they belong to the same chemosyndrome) but maintain
their chemical integrity even when sympatric and are considered distinct
species (Elix 1982).
Neofuscelia parasitica Elix sp. nov. Fig. 11
Species cum thallo ut in Neofuscelia pulla sed ab hac specie acido
glomelliferonico, acido loxodellonico et acido glomellonico continente
differt.
Type: Australia. Australian Capital Territory. On sandstone rocks, Kowen
forest, 16 km east of Canberra, 35°15'S, 149°16'E, 730 m, J. A. Elix 1329,
21.xi.1975; CBG-holotype.
Thallus small foliose, growing over thalli of Neofuscelia verisidiosa
(Essl.) Essl. or Neofuscelia loxodella (Essl.) Essl., moderately adnate, olive-
brown to reddish or dark brown, often paler at the periphery, to 2.0 cm
diam. Lobes irregular to sublinear-elongate, 0.8-1.5 mm wide, contiguous to
imbricate in part. Upper surface dull to slightly shiny at the apices, smooth,
lacking soredia and isidia. Medulla white. Lower surface black, sometimes
dark brown towards the apices, moderately to densely rhizinate, rhizines
slender, simple, black. Pycnidia and apothecia not seen.
Chemistry. Cortex K-, HNO3+ dark blue-green, medulla K-, C-, KC+ rose, P-;
containing glomelliferonic acid (major), loxodellonic acid (major), and
glomellonic acid (trace).
iy rd
This unique new species could be confused with Neofuscelia delisea
(Duby) Essl. since both species lack isidia, have a black lower surface, and
give comparable medullary spot tests. However, N. delisea contains
glomelliferic acid, loxodellic acid and glomellic acid whereas N. parasitica
contians the isostructural depsidones, glomelliferonic acid, loxodellonic acid
and glomellonic acid. This chemistry has been observed previously in N.
subincerta (Essl.) Essl. (an isidiate species) and N. pyrenaica (Essl.) Essl. (Elix,
Jenkins & Jenie 1987). The latter species differs from N. parasitica in the
much larger thalli (2-9 cm diam.), pale brown to black-brown lower surface
and saxicolous substrate. At present N. parasitica has only been found
growing over thalli of the common isidiate Neofuscelia species, N.
verisidiosa and N. loxodella. Interestingly the former species produces
medullary divaricatic acid, a depside unrelated structurally to the
depsidones present in N. parasitica. At present this new species is only
known from the type locality where it appears quite common.
Specimens Examined
AUSTRALIAN CAPITAL TERRITORY. Type locality, J. A. Elix 921,
13.vi.1975, J. A. Elix 21807A, 25.x.1987 (ANUC).
Neofuscelia scabrosina Elix sp. nov. Fig. 12
Species cum thallo ut in Neofuscelia luteonotata sed ab hac specie
acido constipatico, acido protoconstipatico, scabrosini 4,4'-diacetate,
scabrosini 4-acetate-4'- sputyzate, scabrosini 4,4'-dibutyrate et acido ursolico
continente differt.
Type: Australia. Western Australia. On soil and termite mounds at base of
rock outcrop, Camels Peak, N. Sammy, 24.v.1972 ; holo-PERTH 810191.
Thallus foliose, terricolous, moderately adnate to the substrate, olive-
brown or dark brown to black-brown, to 8 cm diam. Lobes irregular to
sublinear-elongate, 1.0-3.0 mm wide, discrete to imbricate in part. Upper
surface dull to slightly shiny and smooth at the apices but becoming
markedly rugose within the thallus, lacking soredia and isidia. Medulla
white. Lower surface pale tan to light brown, sometimes darkening towards
the lobe apices, moderately to densely rhizinate, rhizines short, simple,
concolorous with the lower surface or darkening. Pycnidia common,
conidia bifusiform, sometimes only weakly so, 6-8 x 1 um. Apothecia
common, short stipitate, 3-5 mm in diam., disc black-brown, concave at first
but becoming more or less flat or undulating, margin thin, entire; spores (8),
colourless, ellipsoid, 6-9 x 3.5-5.5 um.
Chemistry. Cortex K-, HNO+ dark blue-green, medulla K-, C-, P-; containing
constipatic acid (major), protoconstipatic acid (major), scabrosin 4,4'-
113
‘ee
3
=
B:
Figures 9-12. New species of Parmeliaceae: 9, Neofuscelia kondininensis
(holotype in CBG); 10, Neofuscelia imatatricoides (holotype in CBG), 11,
Neofuscelia parasitica (holotype in CBG); 12, Neofuscelia scabrosina
(holotype in PERTH). Scale bar = 5 mm.
114
diaceatate, scabrosin 4,4'-dibutyrate, scabrosin 4-acetate-4'-butyrate and
ursolic acid.
This new species could be confused with Neofuscelia luteonotata (J.
Stein.) Essl. since both species often occur on soil, have similar foliose thalli
with a pale tan lower surface, and give completely negative medullary spot
tests. The two species have slightly different general appearance which is
difficult to define and positive identification must rely on chemical analysis.
Thus N. scabrosina contains the fatty acids constipatic acid and
protoconstipatic acid and three scabrosin derivatives while N. luteonotata
contains stenosporic acid and/or divaricatic acid, sometimes with accessory
gyrophoric acid. Furthermore N. scabrosina has a very limited distribution
(Western Australia) while N. luteonotata is widely distributed in southern
Europe, north and south Africa as well as Australia. Given the divergent
distributions and unrelated chemistry these two taxa are considered distinct
species (Elix 1982). Chemically this new species most closely resembles N.
scabrella (Essl.) Essl. from South Africa, but the latter species lacks the fatty
acids, and is distinguished morphologically by the elongate, dichotomously
branched, loosely entangled lobes. At present this new species is only
known from the type locality.
Specimen Examined
WESTERN AUSTRALIA. Type locality, N. Sammy, 24.v.1972 (PERTH).
Neofuscelia subbarbatica Elix sp. nov. Fig. 13
Species cum thallo ut in Neofuscelia parviloba sed ab hac specie
thallo crassiore et acido barbatico, acido 4-O-demethylbarbatico et acido
norobtusatico continente differt.
Type: Australia. Western Australia. On low granite rocks in sclerophyll
forest, 40 km south-east of Armadale, beside the Albany highway, G.
Rambold 5276, 1.iv.1986 ; holo-M.
Thallus small foliose or becoming subcrustose in the centre, tightly
adnate, dark brown, 1-4 cm in diam. but coalescing into larger patches.
Lobes sublinear to irregular, 0.2-0.8 mm wide. Upper surface more or less
smooth at the periphery, convex within and becoming fissured and rugose
to subareolate, dull throughout or slightly shiny at the apices, without
isidia. Medulla white. Lower surface pale tan to pale brown, smooth and
more or less dull, in part attached directly to the substrate, elsewhere
sparsely rhizinate, the rhizines concolorous or darkening, small and fine, to
0.2 mm long. Pycnidia common, conidia weakly bifusiform, 5-6 x 1 um.
Apothecia common, sessile or short stipitate, concave at first but becoming
more or less flat or undulating, to 0.5-2.0 mm in diam., margin entire;
spores (8), colourless, ellipsoid, 7-9 x 4-6 um.
115
Chemistry. Cortex K-, HNO3+ dark blue-green; medulla K-, C-, KC+ yellow,
P-; containing barbatic acid (major), norobtusatic acid (minor), 4-O-
demethylbarbatic acid (trace).
Morphologically this new species closely resembles Neofuscelia
parviloba (Essl.) Essl. in the diminutive subcrustose thalli, the pale lower
surface and the absence of isidia. However N. parviloba has a flatter,
thinner thallus and contains fumarprotocetraric acid and protocetraric acid.
N. melanobarbatica (Essl.) Essl. the only other barbatic acid containing
Neofuscelia with a pale lower surface is distinctly foliose and loosely adnate.
At present this new species is only known from the type collection.
Paraparmelia yamblaensis Elix sp. nov. Fig. 14
Species cum thallo ut in Paraparmelia lithophiloides sed ab hac specie
thallo laxe adnato, lobi latioribus et acido norstictico et acido salazinico
noncontinente differt.
Type: Australia. New South Wales. On rocks in grassland, Great Yambla
Ridge, 17 km south-south-east of Culcairn, 35°50'S, 147°04'E, 580 m, J. A. Elix
23095, 16.xi.1989; holo: CBG, iso: MEL.
Thallus foliose, loosely adnate on rocks, pale mineral grey but
darkening with age, irregularly lobate, 6-8 cm in diam. Lobes variable,
irregular to sublinear-elongate, moderately to sparingly imbricate, 2.0-3.0 (-
5.0)mm wide, the apices crenulate, more or less rotund, occasionally
developing narrower (0.5-1.5 mm), subdichotomously divided secondary
lobes. Upper surface dull, emaculate, lacking soredia and isidia, darkening
with age, the margins commonly blackened. Medulla white. Lower surface
jet black except for a narrow brown marginal zone, very sparsely rhizinate,
rhizines concolorous with the lower surface, simple, short, slender.
Pycnidia rare; conidia bifusiform, 6-9 x 1 um. Apothecia scattered, adnate, 3-
8 mm in diam.; disc concave then undulate-distorted, brown to black-
brown, margin thick and involute at first, then thin, persistent; spores (8),
colourless, ellipsoid, 7-8 x 5-6 um.
Chemistry: Cortex K+ yellow; medulla K-, C-, KC-,.P-; containing
atranorin.
This new species is characterised by the large, loosely adnate thalli,
the broad sublinear lobes with a black lower surface which lack isidia, and
the absence of medullary chemistry. The more adnate portions of the
thallus resemble P. lithophiloides (Kurok.) Elix & Johnston, but that species
has narrower lobes (1-2 mm wide), denser rhizines and contains the
medullary depsidones, norstictic acid and salazinic acid. P. yamblaensis is
only known from the type locality where it is particularly common.
116
Specimens Examined
NEW SOUTH WALES. Type locality, J. A. Elix 23106, 16.xi1.1989 (ASU, PRE);
Ju A, ElpeZ3T 22h SAGX 23123 MiG xt 989K ANDS)
Parmelia crowii Elix sp. nov. Fig. 15
Species cum thallo ut in Parmelia erumpens sed ab hac specie isidiis
coralloidibus et esorediatis differt.
Type: Australia. New South Wales. Brigadoon Farm, Bunga, 18 km south
of Bermagui, 36°34'S, 150°03'E, 30 m, J. A. Elix 23373, 2.xii.1989; holo: CBG;
iso ANUC, HO. MEL, PRE.
Thallus saxicolous, adnate to loosely attached, pale mineral-grey, 8-20
cm in diam. Lobes short, subirregular to apically rotund, imbricate, 2-5 mm
wide. Upper surface shiny, plane, white-reticulate at first but becoming
conspicuously cracked to the margin, pseudocyphellae effigurate, somewhat
raised, dense, fusing into a reticulate network over the whole surface,
isidiate; the isidia epicorticate, coarse, cylindrical at first then coralloid,
rarely bursting apically and not becoming granular and sorediose. Lower
surface moderately rhizinate, rhizines simple or squarrosely branched, 1-2
mm long. Pycnidia and apothecia not seen.
Chemistry. Cortex K+ yellow; medulla K+ yellow then deep red, C-, P+ red-
orange; containing atranorin, chloroatranorin, salazinic acid (major),
consalazinic acid (minor), norstictic acid (trace).
This new species closely resembles P. erumpens Kurok. except in the
develpment of asexual propagules. Thus P. erumpens is sorediate-isidiate
where the coarse granular soredia become in part corticate, forming dense
marginal and laminal soralia and/or extended, granular, coralloid-isidioid
outgrowths while in P. crowii the coarse cylindrical isidia become densely
coralloid branched, but only very rarely burst open and then do not become
sorediate. Parmelia pseudotenuirima Gyelnik, another Australian species
with cylindrical isidia, differs in having much thinner (less than 0.05 mm
vs. 0.2-0.3 mm thick), syncorticate isidia. This new species is named in
honour of Dr. W. D. Crow, friend, colleague and owner of Brigadoon Farm.
Parmelina euplectina Kurok. ex. Elix sp. nov. Fig. 16
Species cum thallo ut in Parmelina pseudorelicina sed ab hac specie
medulla inferiore pigmento ochraceo et euplectino continente differt.
117
Figures 13-16. New species of Parmeliaceae: 13, Neofuscelia subbarbatica
(holotype in M); 14, Paraparmelia yamblaensis (holotype in CBG); 15,
Parmelia crowii (holotype in CBG); 16, Parmelina euplectina (holotype in
MEL). Scale bar = 5 mm.
118
Type: Australia. New South Wales. On Casuarina species, Raymond
Terrace to Buladelah road, 30 km north of Karuah, R. Filson 7176, 9.v.1965;
holo: MEL.
Thallus corticolous, adnate to tightly adnate, pale grey to grey-green,
2-4 cm diam. Lobes sublinear-elongate, generally narrow but occasionally
more robust, 1.0-3.0 mm wide, irregularly branched, imbricate, margins
sinuous, shining, black, ciliate, the cilia sparse, black, simple, short, 0.1-0.3
mm long, mainly in the axils of the lobes. Upper surface shiny, plane or
weakly foveolate at first, becoming rugulose with age, emaculate, lacking
soredia and isidia. Upper medulla white, lower medulla orange-yellow.
Lower surface black, with a narrow, brown, naked marginal Zone,
moderately to sparsely rhizinate, rhizines simple, black. Pycnidia
numerous, conidia cylindrical to weakly fusiform, 6-7 x 0.7 um. Apothecia
common, sessile to substipitate, 1-2.5 mm diam., disc concave, margin thin,
entire, not crenulate or inflexed, thalline exciple smooth; spores (8),
colourless, ellipsoid, 9-13 x 6-7 um.
Chemistry. Cortex K+ yellow, medulla K-, C+ red, P-, lower medulla K+
purple; containing atranorin, chloroatranorin, lecanoric acid (major),
euplectin.
Although this new species closely resembles Parmelina
pseudorelicina (Jatta) Kantvilas & Elix, it is clearly distinguished by the
partially orange-yellow pigmented lower medulla due to the presence of the
anthraquinone, euplectin. To date this is the only species of Parmelina
known with a pigmented medulla. This species was formerly referred to as
Parmelia jejunga Kurok. but the latter is a nomen nudum (Filson 1983). P.
euplectina is a very rare endemic species, currently known only from the
type locality.
Parmelina johnstoniae Elix sp. nov. Fig..17
Species cum thallo ut in Parmelina conlabrosa sed ab hac specie thallo
brunneo-cinerascens, fragiliore et acido protolichesterinico et acido
lichesterinico continente differt.
Type: Australia. New South Wales. On trunk of Acacia, Buckenbowra River
Estuary, 7.5 km W of Batemans Bay, 34°42'S, 150°06'E, 5 m, J. Johnston 2610;
holo: CBG.
Thallus corticolous or lignicolous, adnate, brownish-grey to pale
mineral grey, 2-6 cm in diam. Lobes variable, sublinear-elongate to
subirregular and apically subrotund, 1-3 mm wide, sometimes
subdichotomously branched, marginal cilia moderate to sparse and
concentrated in the lobe axils, simple. Upper surface shiny, emaculate,
more or less plane, moderately to densely isidiate, the isidia cylindrical,
119
erect, often branched, less than 0.5 mm high. Medulla white. Lower
surface black, moderately rhizinate, the rhizines black, shiny simple.
Pycnidia and apothecia not seen.
Chemistry. Cortex K+ yellow, medulla K-, C-, KC-, P-; containing atranorin,
chloroatranorin, lichesterinic acid, protolichesterinic acid.
This new species is distinguished by the cylidrical isidia and
medullary chemistry. Morphologically this species closely resembles the
common P. conlabrosa (Hale) Elix & Johnston but differs in the browner,
more fragile thallus and the negative medullary reactions. P. conlabrosa
contains medullary lecanoric acid. P. johnstoniae is probably the isidiate
counterpart of P. endoleuca (Taylor) Hale. At present this new species is
only known from the type locality where it appears quite common. This
species is named in honour of my former colleague, Ms. Jen Johnston.
Specimens Examined
NEW SOUTH WALES. Type locality, J. A. Elix 10978, 29.v.1983, J. A. Elix
22618, 4.vili.1988 (ANUC).
Parmelinopsis protocetrarica Elix sp. nov. Fig. 18
Species cum thallo ut in Parmelinopsis minarum sed ab hac specie
subtus pallido et acido protocetrarico continente differt.
Type: Australia. Queensland. On mossy trunk of felled tree in rainforest,
Walter Hill Range, 26 km south-east of Ravenshoe, 17°46'S, 145°41'E, 800 m,
J. A. Elix 17041 & H. Streimann, 2.vii.1984; holo: CBG.
Thallus corticolous, adnate, whitish to pale greenish mineral grey, 2-4
cm diam. Lobes sublinear-elongate, contiguous, 0.8-1.2 mm wide, the
marginal cilia irregularly dispersed, simple, to 0.7 mm long. Upper surface
shiny, sparingly maculate at the apices, plane, densely isidiate, the isidia
cylindrical, erect, often branched, to 0.5 mm high, becoming procumbent
and flattened with age. Medulla white. Lower surface ivory to pale brown,
black-brown at the apices, moderately rhizinate, the rhizines black, shiny,
simple or squarrose branched. Pycnidia and apothecia not seen.
Chemistry. Cortex K+ yellow medulla K-, C+ rose, KC+ red, P+ orange;
containing atranorin, gyrophoric acid (minor), protocetraric acid (major),
unknown (minor).
Morphologically this new species superficially resembles
Parmelinopsis minarum (Vainio) Elix & Hale with fragile thalli with
densely isidiate narrow lobes. However P. protocetrarica is clearly
distinguished by the pale lower surface of the lobes (black in P. minarum)
120
and the presence of additional protocetraric acid in the medulla. At present
this new species is only known from the type collection.
Parmotrema submerrillii Elix sp. nov. Fig. 19
Species cum thallo ut in Parmotrema merrillii sed ab hac specie lobis
sine laciniis marginalibus differt.
Type: Australia. Queensland. On tree in picnic area adjacent to rainforest,
Mt. Spec National Park, ridge on the Loop, on the Paluma road, WNW of
Townsville, 1000 m, M. E. Hale 64052, 28.vii.1983; holo: CBG.
Thallus corticolous, loosely adnate, coriaceous, pale mineral grey or
darkening with age, 8-10 cm diam. Lobes irregular, 5-15 mm wide, with
rotund apices, margins ciliate, cilia prominent, 1.0-3.5 mm long. Upper
surface emaculate, lacking soredia and isidia. Medulla white. Lower surface
black, marginal rim brown, sparsely rhizinate, rhizines simple, black.
Pycnidia common, conidia sublageniform, 5-6 x 1 um. Apothecia common,
stipitate, 4-8 mm diam., imperforate, the exciple dentate. ciliate,
amphithecium rugose, white maculate; spores (8), colourless, ellipsoid, 25-
34 x 12-16 um.
Chemistry. Cortex K+ yellow, medulla K-, C-, KC-, P+orange-red; containing
atranorin, chloroatranorin, protocetraric acid (major).
This new species resembles the south-east Asian species, P. merrillii
(Vainio) Hale both chemically and morphologically. However where P.
merrillii develops long narrow laciniae on the lobes and frequently on the
exciple of the apothecia, these are absent in P. submerrillii, although the
exciple is dentate in the latter. This species may also be confused with P.
zollingeri (Hepp) Hale as they have similar thalli and contain protocetraric
acid. However P. zollingeri has very sparse, short (less than 1 mm) cilia,
smaller spores (18-22 x 8-10 um) and an eciliate exciple. At present this new
species is only known from the type locality.
Xanthoparmelia austrocostrictans Elix sp. nov. Fig. 20
Species cum thallo ut in Xanthoparmelia constrictans sed ab hac
specie superne emaculata et rhizinis modicis differt.
Type: Australia. New South Wales. On sandstone rocks in open Eucalyptus
woodland, Morton National Park, 8 km north-east of Nerriga, J.A. Elix 3070;
CBG-holotype.
Thallus foliose, moderately to loosely adnate to rocks or soil, yellow-
green in colour darkening with age, 5-7 cm in diam. Lobes linear-
elongate, dichotomously to subdichotomously branched, moderately to
121
Figures 17-20. New species of Parmeliaceae: 17, Parmelina johnstoniae (J. A.
Elix 10978 in ANUC); 18, Parmelinopsis protocetrarica (holotype in CBG); 19,
Parmotrema submerrillii (holotype in CBG); 20, Xanthoparmelia
austroconstrictans (holotype in CBG). Scale bar = 5 mm
122
sparingly imbricate, 0.4-1.5 mm wide but often irregularly widened and
constricted. Upper surface smooth, shiny, emaculate, lacking soredia and
isidia but with numerous pycnidia, margins black; medulla white. Lower
surface jet black but brown at the tips of the lobes, moderately rhizinate,
rhizines simple or tufted. Pycnidia common, conidia bifusiform, 5-8 x 0.7
um. Apothecia not seen.
Chemistry. Cortex K-, medulla K+ yellow becoming deep red, C-, P+ orange;
containing usnic acid, salazinic acid and consalazinic acid.
X. austroconstrictans is distinguished by the loosely imbricate, linear
elongate lobes which are irregularly constricted and the production of
salazinic acid in the medulla. Chemically X. austroconstrictans is identical
with X. rubrireagens - a morphologically similar species. However the lobes
of X. rubrireagens tend to be much more uniform in width (i.e. not
irregularly constricted) and the underside is sparsely rhizinate and often
erhizinate towards the apices, the lower surface rugose and wrinkled, and
the rhizines themselves short, robust and + tufted. By contrast X.
austroconstrictans is moderately rhizinate, with + subapically grouped
rhizines and the lower surface is more or less plane. This new species also
resembles the South African X. constrictans (Nyl.) Hale - a species which
differs in having a maculate upper surface and dense, long, + branched,
curved rhizines commonly projecting beyond the margins of the lobes. X.
austroconstrictans is a rare species in Australia, being restricted to the
mountains of south-eastern Australia (New South Wales and Victoria) and
Tasmania.
Specimen Examined
VICTORIA. Bullock Creek, 16 km south of Bendigo along Calder Highway, J.
Johnston 1162A (ANUC). TASMANIA. Royal George Mine area, G.C. Bratt
345 (HO).
Xanthoparmelia canobolasensis Elix sp. nov. Fig. 21
Species cum thallo ut in Xanthoparmelia glareosa sed ab hac specie
thallo sparsim rhizinato, rhizinis simplicibus et acido hypostictico et acido
hyposalazinico continente differt.
Type: Australia. New South Wales. On volcanic rock ledge in open
Leptospemum scrub, west slope of Mt. Canobolas, 8 km south-west of
Orange, 32°21'S, 148°59'E, 1300 m, J. Johnston 2909, 6.xii.1989; holo: CBG.
Thallus foliose, adnate to tightly adnate, saxicolous or terricolous,
yellow-green but becoming greyish with age, 5-10 cm in diam. Lobes
subirregular, elongate, imbricate, 1.0-3.0 mm wide, with distinct black
margins; secondary lobes 0.4-1.0 mm wide, subdichotomously branched,
123
overlying the centre of the thallus but sometimes becoming dominant.
Upper surface dull or commonly shiny, emaculate, more or less smooth,
lacking soredia and isidia; medulla white. Lower surface black, brown-black
at the apices, sparsely rhizinate, rhizines simple, concolorous with the lower
surface. Pycnidia common, conidia bifusiform 5-7 x 1 um. Apothecia
sessile, 2.0-4.0 mm in diam., concave; disc cinnamon-brown, margin thin,
strongly involute at first but becoming crenulate and lacerate; spores (8),
colourless, ellipsoid, 7-10 x 5-6 um.
Chemistry. Cortex K-; medulla K+ yellow becoming red, C-, KC-, P+ yellow-
orange; containing usnic acid, norstictic acid (major), connorstictic acid
(minor), hypostictic acid (minor/trace), and hyposalazinic acid (trace).
This new species resembles X. glareosa (Kurok. & Filson) Elix &
Johnston in growth form and chemistry, but X. glareosa is moderately
rhizinate with simple and dichotomously branched rhizines and contains
medullary salazinic acid in addition to norstictic and connorstictic acids
whereas X. canobolasensis is sparsely rhizinate, has simple rhizines and
contains hypostictic and hyposalazinic acids in addition to norstictic and
connorstictic acids.
Specimen Examined
NEW SOUTH WALES. On soil in grassy area, Orange View, Pinnacles Road,
Mt. Canobolas, 1100 m, M. E. Hale 59947, 25.11.1984 (ANUC).
Xanthoparmelia hyposalazinica Elix sp. nov. Fig. 22
Species cum thallo ut in Xanthoparmelia subnuda sed ab hac specie
lobis sublineariter elongatis et acidum hyposalazinicum continente differt.
Type: Australia. Australian Capital Territory. On porphyry rocks in pasture,
0.8 km west of Coppins Crossing, 8 km west of Canberra, 35°17'S, 149°03'E,
520 my fA Elix Al/753,17 41992) holo: CBG:
Thallus foliose, loosely adnate on rocks, yellowish-green, 3-5 cm in
diam. Lobes sublinear-elongate, di- or trichotomously branched, imbricate,
often ascending at the apices, 0.5-1.5 mm wide. Upper surface smooth,
glossy or opaque, emaculate, often blackened at the margins, lacking soredia
and isidia; medulla white. Lower surface black, nude or sparsely rhizinate,
rugulose, rhizines simple, black, robust, 0.2-1.0 mm long. Pycnidia
common; conidia bifusiform, 5-7 x 1 um. Apothecia sessile to substipitate, 2-
4 mm in diam., margins undulate or not, disc dark brown, thalline exciple
smooth, emaculate; spores (8), colourless, ellipsoid, 9-12 x 5-7 um.
Chemistry. Cortex K-, medulla K+ yellow then red, C-,KC-, P+ yellow;
containing usnic acid and hyposalazinic acid.
124
Its narrow, sublinear-elongate lobes, lack of maculae, soredia and
isidia together with the black lower surface and medullary hyposalazinic
acid distinguish this species. X. subnuda (Kurok.) Elix, a related species, has
more linear-elongate lobes and contains medullary norstictic acid and
connorstictic acid. Currently this rare endemic species is only known from
the type locality.
Xanthoparmelia masonii Elix sp. nov. BigaZ3
Species cum thallo ut in Xanthoparmelia mougeotina sed ab hac
specie thallo subtus brunneo differt.
Type: Australia. New South Wales. On granite outcrops in pasture with
Eucalyptus, M. E. Hale 59,276, 10.i.1982; holo: CBG.
Thallus small foliose to subcrustose, tightly appressed on rocks,
greenish-yellow in colour, subirregularly lobate or becoming almost
subcrustose towards the centre, 3-6 cm in diam. Lobes sublinear, discrete or
contiguous, 0.3-1.0 mm wide, older lobes irregularly fractured and areolae
commonly formed. Upper surface opaque, emaculate, often tangentially
cracked, moderately to sparsely isidiate, the isidia cylindrical, mainly simple;
medulla white. Lower surface ivory to pale brown but often straw-yellow
near the apices of the lobes, moderately rhizinate, the rhizines simple, pale
brown. Pycnidia and apothecia not seen.
Chemistry. Cortex K-, medulla K+ yellow, C-, KC-, P+ orange; containing
usnic acid, stictic acid, constictic acid, norstictic acid (minor), cryptostictic
acid (trace), rarely menegazziaic acid ( trace), and associated unknown (+
trace).
Morphologically this species closely resembles X. mougeotina (Nyl.)
D. Gall. in that both lichens have tightly adnate thalli, contiguous lobes,
cylindrical isidia and medullary stictic acid. However X. mougeotina is
readily distinguished by the black lower surface. Another related species, X.
victoriana Elix & Johnston has a pale lower surface, but is distinguished by
the inflated, erumpent isidia. At present this new species is known only
from the type collection.
Xanthoparmelia trirosea Elix sp. nov. Fig. 24
Species cum thallo ut in Xanthoparmelia multipartita sed ab hac
specie lobi lineariter angustatis, modice imbricatis et acido hypoconstictico
adjectus continente differt.
125
Figures 21-24. New species of Parmeliaceae: 21, Xanthoparmelia
canobolasensis (holotype in CBG); 22, Xanthoparmelia hyposalazinica
(holotype in CBG); 23, Xanthoparmelia masonii (holotype in CBG); 24,
Xanthoparmelia trirosea (isotype in ANUC). Scale bar (Figs. 21, 22, 24) =5
mm; (Fig. 23) = 2 mm.
126
Type: Australia. Australian Capital Territory. On granite rocks in open
woodland, Gudgenby River Gorge, 4.5 km south of Tharwa, 35°34'S,
149°04'E, 620 m, J. A. Elix 10165, 12.iv.1982; hoio: CBG, iso: ANUC, MEL JJ. A.
Elix : Lichenes Australasici Exsiccati, No. 19, distributed as Parmelia
multipartita (R. Br. ex. Crombie) Elix].
Thallus foliose, very loosely adnate on rocks, yellow to yellow-green
but blackening with age, 6-9 cm in diam. Lobes _ linear-elongate,
subdichotomously branched, divaricate in places, moderately imbricate,
becoming suberect and exposing the black lower surface, 0.5-1.0 mm wide.
Upper surface smooth, shiny, emaculate, lacking soredia and isidia but with
black margins; medulla white. Lower suface jet black, wrinkled, with very
sparse rhizines or rhizines absent, rhizines simple, short, robust. Pycnidia
common; conidia bifusiform, 5-7 x 1 um. Apothecia adnate, 1.0-3.0 (-10.0)
mm in diam., disc chestnut brown at first but darkening with age, margin
entire, thick and involute at first but becoming thin and subentire with age;
spores (8), colourless, ellipsoid, 6-10 x 3.5-5.5 um.
Chemistry. Cortex K-, medulla K+, pale reddish, C-, KC-, P-; containing
usnic acid, hypostictic acid, hyposalazinic acid, hypoconstictic acid .
This species has previously been confused with Xanthoparmelia
multipartita (Crombie) Hale, but these two taxa show consistent
morphological and chemical differences. xX. multipartita differs
morphologically in having more densely imbricate lobes and marginal lobes
with a brown lower surface, whereas the thalli of X. trirosea are only
moderately imbricate and the lower surface is jet black to the margins.
Chemically X. trirosea contains hypoconstictic acid in addition to hypostictic
acid and hyposalazinic acid observed in X. multipartita. The name of this
species refers to the three rose-red spots which develop on a TLC plate after
spraying with 10% sulfuric acid and charring. The latter species also
contains minor or trace amounts of 4-O-methylhypoprotocetraric acid
which is absent in X. trirosea. This new species occurs on exposed, acidic
rocks in areas of moderate rainfall in coastal and hinterland areas of south-
eastern Australia (N.S.W., A.C.T., Tas.).
Specimens Examined
NEW SOUTH WALES: On granite rocks, 10 km N of Jindabyne along the
Cooma road, J. A. Elix 1653, 1654, 1818, 21.1.1976 (ANUC). TASMANIA: On
granite rocks along the foreshore, Bicheno, J. A. Elix 5531, 13.1.1979 (ANUC).
New Combinations and Name
Bulbothrix queenslandica (Elix & Stevens) Elix, comb. nov.
Basionym: Parmelia queenslandica Elix & Stevens, Aust. J. Bot. 27: 873
(1979).
127,
Bulbothrix subtabacina (Elix) Elix, comb. nov.
Basionym: Parmelia subtabacina Elix, in Elix & Stevens, Aust. J. Bot. 27:
875 (1979).
Canoparmelia owariensis (Asah.) Elix, comb. nov.
Basionym: Parmelia owariensis Asah., J. Jap. Bot. 28: 135 (1953).
Canoparmelia pustulescens (Kurok.) Elix, comb. nov.
Basionym: Parmelia pustulescens Kurok. in Hale & Kurok., Contrib.
U.S. Nat. Herb. 36: 156 (1964).
Canomaculina melanochaeta (Kurok.) Elix, comb. nov.
Basionym: Parmelia melanochaeta Kurok. in Hale & Kurok., Contrib.
U.S. Nat. Herb. 36: 133 (1964).
Imshaugia evernica (Elix & Johnst.) Elix, comb. nov.
Basionym: Parmeliopsis evernica Elix & Johnst., Mycotaxon 31: 495
(1988).
Parmotrema pseudovirens (Gyelnik) Elix, comb. nov.
Basionym: Parmelia pseudovirens Gyelnik, Fedde, Repert. Spec.
Nov. 29: 288 (1931).
Rimeliella haitiensis (Hale) Elix, comb. nov.
Basionym: Parmelia haitiensis Hale, Bryologist, 62: 20 (1959).
Xanthoparmelia xanthofarinosa Elix , nom. nov.
Thallus adnatus vel arcte adnatus, saxicola, flavovirens, 3-5 cm diametro;
lobi sublineares vel subirregulares, 0.5-2.0 mm lati. Superfices nitida,
emaculata, dense sorediata, soraliis globoso-capitatis, confluentibus, et
sorediis farinosis; superfices inferior castanea sed apices loborum versus
saepe atrobrunnescens, modice rhizinata. Thallus acidum usnicum,
acidum constipaticum, acidum_ protoconstipaticum, loxodinum,
conloxodinum, norlobaridonum, conorlobaridonum et substantia ignota
continens.
Type: Australia. Australian Capital Territory. On sandstone rocks, Kowen
Forest, 16 km east of Canberra, 730 m, J. A. Elix 1830; holo: MEL; iso: CBG.
= Parmelia xanthosorediata Elix (nomen nudum, Chester & Elix 1979).
128
= Xanthoparmelia xanthosorediata (Elix) Elix & Johnston, Bull. Br.
Mus. nat. Hist. (Bot.) 15: 357 (1986) (nomen nudum, Begg, Chester &
Elix 1979).
ACKNOWLEDGEMENTS
I thank the Australian Research Council for generous financial support
of this project and the late Dr. M. E. Hale for his cooperation, generous
advice and assistance in obtaining critical type material in the early part of
this project. I also wish to thank the staff of the Science Photographic
Unit, ANU, for preparing the photographs, Mr. D. Verdon for checking
the Latin descriptions and Ms. C. E. Barclay and S. A. Elix who determined
the chemistry of many specimens. I thank the following colleagues and
herbaria for making collections available: Dr. R. B. Filson (Booral), Ms. J.
Johnston (ANUC), Dr. G. Kantvilas (HO), Dr. P. M. McCarthy (MEL), Mr.
N. Sammy (PERTH), Dr. G. Rambold (M) and Mr. J. Whinray (MEL).
LITERATURE CITED
Begg, W. Rv Chester,’ D.’ Orie PEiix, J. A: (1979).. The structure of
conorlobaridone and conloxodin. New depsidones from the lichen
Xanthoparmelia xanthosorediata. Aust. J. Chem. 32: 927-929.
Chester, D. O. & Elix, J. A. (1979). Three new aliphatic acids from lichens of
genus Parmelia (subgenus Xanthoparmelia). Aust. J]. Chem. 32: 2565-
2569.
Culberson, C. F. (1972). Improved conditions and new data for the
identification of lichen products by a standardized thin-layer
chromatographic method. J. Chromatogr. 72: 113-125.
Culberson, C. F.. Culberson, W. L. and Johnson, A. (1981). A standardized
TLC analysis of B-orcinol depsidones. Bryologist 84: 16-29.
Culberson, C. F. and Johnson, A. (1982). Substitution of methyl tert.-butyl
ether for diethyl ether in the standardized thin-layer chromatographic
method for lichen products. J. Chromatogr. 238: 483-487.
Elix, J. A. (1982). Peculiarities of the Australasian lichen flora: accessory
metabolites, chemical and hybrid strains. J. Hattori Bot. Lab., 52: 407-
415.
Elix, J. A., Jenie, U. A. & Jenkins, G. A. (1987). Three new depsidones from
the lichen Neofuscelia subincerta. Aust. J. Chem. 40: 2031-2036.
Elix, J. A. & Johnston, J. (1986). New Species of Relicina (Lichenized
Ascomycotina) from Australasia. Mycotaxon, 27: 611-616.
129
Elix, J. A. & Johnston, J. (1987a). New Species of Paraparmelia (Lichenised
Ascomycotina) from Australia and New Zealand. Brunonia 1986, 9:
139-193.
Elix, J. A. & Johnston, J. (1987b). New species of Parmelina (Lichenised
Ascomycotina) from Australia and New Zealand. Brunonia 1986, 9:
155-161.
Elix, J. A. & Johnston, J. (1987c). New species and new records of
Xanthoparmelia (Lichenized Ascomycotina) from Australia.
Mycotaxon 29: 359-372.
Elix, J. A. & Johnston, J. (1988a). New species in the lichen family
Parmeliaceae (Ascomycotina) from the Southern Hemisphere.
Mycotaxon 31: 491-510.
Elix, J. A. & Johnston, J. (1988b). New species in the lichen genus
Paraparmelia (Ascomycotina) from the Southern Hemisphere,
Mycotaxon 32: 399-414.
Elix, J. A. & Johnston, J. (1988c). New Species and new reports of
Flavoparmelia (lichenized Ascomycotina) from the Southern
Hemisphere, Mycotaxon 33: 391-400.
Elix, J. A., Johnston, J. & Armstrong, P. M. (1986a). A revision of the lichen
genus Xanthoparmelia in Australasia. Bull. Br. Mus. nat. Hist. (Bot.)
15: 163-362.
Elix, J. A., Johnston, J. and Parker, J. L. (1987). A Catalogue of Standardized
Thin Layer Chromatographic Data and Biosynthetic Relationships for
Lichen Substances (Aust. Nat. University, Canberra).
Elix, J. A., Johnston, J. and Parker, J. L. (1988). A Computer Program for the
Rapid Identification of Lichen Substances. Mycotaxon 31: 89-99.
Filson, R. B. (1983). Checklist of Australian Lichens, Ist Edn. National
Herbarium of Victoria, Public Lands and Forests Division, Department
of Conservation, Forests and Lands, p 70.
Lumbsch, H. T. and Elix, J. A. (1985). A new species of the lichen genus
Diploschistes from Australia. Pl. Syst. Evol., 150: 275-279.
Nash, T. H. III & Elix, J. A. (1987). New species and new reports in the
Parmeliaceae (Lichenized Ascomycotina) from South Africa and
Australia. Mycotaxon, 29: 467-476.
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MYCOTAXON
Volume XLVII, pp. 131-146 April-June 1993
TAXONOMIC STUDIES IN THE GENUS MYCOSPHAERELLA. 2.
NOTES ON SOME ADDITIONAL SPECIES OCCURRING
ON BRASSICACEAE.
MICHAEL CORLETT
Centre for Land and Biological Resources Research
Agriculture Canada, Research Branch
C.E.F., Ottawa, Ontario, Canada K1A OC6
Abstract
Comments, descriptions and illustrations of additional non Canadian species
of Mycosphaerella occurring on Brassicaceae are provided. Type specimens
of four species were examined: M. denigrans, M. pachyasca, M. pashkiensis
and M. pyrenaica. Several species are only known from their original
descriptions as type or other specimens were unavailable for study or not
located; some species have never been illustrated.
Introduction
A publication (Corlett, 1988) on the species of Mycosphaerella
occurring on Cultivated, wild and weedy members of the family
Brassicaceae in Canada treated four species, M. brassicicola (Duby)
Lindau, M. cruciferarum (Fr.) Lindau, M. densa (E. Rostrup) Lind and
M. tassiana (de Not.) Johans. There are some additional species
described from Brassicaceae by various authors which were not
dealt with in that publication. | have not seen Canadian collections
of these species and they are presently presumed to be extralimital.
some of the names are known to me only from the literature. Type
collections of four species have been examined. | have attempted
to borrow types of several others for examination but they were
132
unavailable. Nevertheless, it was considered worthwhile to discuss
or provide notes or brief descriptions, from personal observations or
from the literature, and where possible to provide illustrations.
Unfortunately for several species known to me only from the
literature and for which types or other collections have not been
examined, there are no published illustrations available.
Species described on Brassicaceae
Mycosphaerella alyssi (Hollés) Moesz, Balkan-Kutat. Tud. Eredm. 3:
132 & 136. 1926.
Sphaerella alyssi Hollés, Ann. Hist.-Nat Mus. Hung. 5: 44.
1907.
This species was described by Hollds (op. cit., 1907, p. 44) on
Alyssum tortuosum Waldst. & Kit. from Hungary. Hollés provided the
following description without illustrations: “Peritheciis gregariis,
epidermide velatis, depresso-globosis, fuscis, punctiformibus,
pertusis, 50-70 p diam., contextu parenchymatico; ascis ovoideis vel
Clavatis, breve st[i]pitatis, octosporis, 24-30 x 9-10 yw aparaphysatis;
sporidiis 2-3 stichis obiongis, medio 1-septatis, non constrictis,
hyalinis, 9-10 x 3-4 yw". Moesz (op. cit., 1926, p. 136), in making the
combination to Mycosphaerella listed Alyssum scardicum Wettst. as
a host from Albania. | only know this species from the literature and
have not seen type or any other collections, nor have | seen any
illustrations of this fungus. M. alyssi was described in Tomilin (1979,
p. 75). He cited the measurements given by Hollés and did not
provide any discussion or illustrations of the asci and ascospores.
Mycosphaerella brassicicola (Duby) Lindau in Engler & Prantl, Die
Natutrlichen Pflanzenf. 1(1): 424. 1897.
Sphaeria brassicicola Duby in de Candolle, Botanicon Gallicum
2: 712. 1830, as brassicaecola, non Sphaeria brassicicola
Berkeley & Broome in Berkeley, Outlines Brit. Fung., p. 401.
1860.
_ Sphaerella brassicicola (Duby) Cesati & de Notaris, Comment.
Soc. Crittog. Ital. 1(3): 238. 1963.
Anamorph: Asteromella brassicae (F. Chevallier) Boerema &
van Kesteren, Persoonia 3: 18. 1964.
133
This important pathogen causing conspicuous ‘ring spot" lesions
on the leaves of Brassica species has been redescribed, illustrated
and discussed by Punithalingam and Holliday (1975) and by Corlett
(1988, p. 63).
Mycosphaerella carniolica (Niessl) Lindau, Hilfsb. Sammeln
Ascomyceten, p. 38. 1903.
Sphaerella carniolica Niessl, Oesterr. Bot. Z. 25(8): 85. 1875;
Hedwigia 14(10): 147. 1875.
Sphaerella carniolica originally was described by Niessl (op. cit.,
1875) from Draba ciliata Scop. without illustrations as follows: "Epi-
rarius hypophylla. Perithecia dense disseminata, minuta, punctiformia,
globosa, papillata, tandem vertice umbilicata, atra; ascis fasciculatis
8sporis, obovatis vel oblongis, sessilibus 30-42 x 14-18; sporidiis
farctis, Ccuneato-oblongis, rectis, medio septatis vix constrictis, dilute
virescentibus 15-19 x 4-5".
This name is known to me only from the literature; | have not
seen the type or any other collections. Tomilin (1979, p. 76)
accepted carniolica as a distinct species of Mycosphaerella but
provided no illustrations. Lind (1934, p. 66) considered Sphaerella
carniolica to be a synonym of Mycosphaerella confinis (Karsten)
Dearness (as confinis (Karsten) Lind). Von Arx (1949, p. 40)
considered both confinis and carniolica to be synonyms of
Mycosphaerella tassiana (de Notaris) Johanson. Niessl’s
measurements for the asci and ascospores of S. carniolica are
rather low for a mature specimen of M. tassiana. | am unable to
comment on the taxonomic status of this species.
Mycosphaerella confinis (P.A. Karsten) Dearness, Rep. Can. Arctic
Exped. 1913-1918 4(C): 6. 1923.
Sphaerella confinis P.A. Karsten, Ofvers. Forh. Kongl. Svenska
Vetensk.-Akad. 1872(2): 106. 1872; Hedwigia 11: 187. 1872.
This species was described by Karsten (op. cit., 1872) from dead
foliage of Braya purpurascens Bunge, Draba martinsiana Gay and D.
wahlenbergii Hartm. from Svalbard, Norway without illustrations as
follows: “Perithecia dense gregaria, amphigena, primitus epidermidi
innata, dein protuberantia, spohaeroidea, interdum vertice obtusissime
134
conoidea, dein ostiolo rotundato hiantia, laevia, glabra, nigra, latit.
circiter 150 mmm. Asci subsessiles, fusoideo-elongati vel elongato-
clavati, vulgo inaequilaterales, longit. 50-65 mmm., crassit. 11-13
mmm. Sporae 8:nae, di- vel subtristichae, aciculari- vel fusoideo-
elongatae, ut plurimum leviter Curvulae, uniseptatae, medio non
constrictae, hyalinae, longit. 16-24 mmm., crassit. 3-5 mmm.
Paraphyses nullae". Measurements given by Saccardo (Syll. F. 1:
507-8. 1882) for S. confinis Karsten differ somewhat from those given
by Karsten and are as follows: perithecia 100-120um, asci 45-52 x
10-13um, ascospores 14-19 x 3-5um. Dearness (op. cit., 1923, p.
6) listed a collection on Anemone parviflora Michx. [Ranunculaceae].
A collection identified as M. confinis in the Dearness herbarium (no.
6018) in DAOM, is on Antennaria margaritacea R. Br. [Asteraceae].
Number 6018 does not agree with Karsten’s description of S.
confinis; the asci of the Dearness specimen are cylindrical and
narrow and measure 45-50 x 6-9um and the ascospores at 14 to
15um in length likewise do not match Karsten’s description.
Von Arx (1949, p. 40) considered Sphaerella confinis Karsten to
be a synonym of Mycosphaerella tassiana (de Notaris) Johanson.
Mycosphaerella cruciferarum (Fries) Lindau in Engler & Prantl, Die
Naturlichen Pflanzenf. 1(1): 324. June 1897.
Sphaeria cruciferarum Fries, Systema Mycol. 2: 525. 1823.
Sphaerella cruciferarum (Fries) Saccardo, Michelia 2: 315.
1881.
A presumably authentic collection of Sphaeria cruciferarum Fr.
from the Fries herbarium at UPS on Cheiranthus L. from France was
seen by me. A microscope slide prepared from this collection
possessed a single fascicle of relatively mature aparaphysate asci,
60-65 x 14-16um, unfortunately without ascospores. While these
asci have the morphology and dimensions of asci of M. cruciferarum
(Corlett, 1988), a positive identification cannot be made. WM.
cruciferarum was recently redescribed and illustrated by Corlett
(1988, p. 64). Despite not having observed type or good authentic
material, this species appears to be well characterized (Barr, 1959,
p. 15; Corlett, op. cit.) by its relatively broad fusoid ascospores and
the presence of a network of dark subepidermal hyphae connecting
individual closely grouped ascomata.
135
Mycosphaerella denigrans (Kirschstein) Tomilin, Opredelitel’gribov
roda Mycosphaerella Johns., p. 77. 1979.
Sphaerella denigrans Kirschstein, Ann. Mycol. 37: 103. 1939.
Mycosphaerella phlomidicola Tomilin, Nov. *Sist. Niz. Rast.
1966, p. 150. 1966. FIG. 1
Sphaerella denigrans was described by Kirschstein (op. cit., 1939,
p. 103) from Erysimum hieraciifolium [given without authority] without
illustrations and described as follows: “Peritheciis dense _ stipatis,
substratum denigratum saepe longe, interdum ex toto inducentibus,
sub peridermio convexo-sublato orientibus, demum plus minus
erumpentibus, ostiolo plano, pertuso, rotundulo instructis, nigris,
globosis, membranaceis, 80-100 yp diam.; contextu fuliginoso, e
cellulis minutis Composito. Ascis aparaphysatis, late cylindraceis
vel clavatis, apice plane rotundatis et incrassatis, vix pedicellatis, 8-
sporis, 25-30 x 7-9 wu. Sporidiis oblique monostichis vel
longitudinaliter di- vel tristichis, hyalinis, medio uniseptatis, oblonge
ellipsoideis, rectis et [subinaequalibus], non constrictis 10-11 x 3 p".
Tomilin (op. cit., 1979, p. 77 & fig. 39) has examined the holotype
collection of Sphaerella denigrans, ex herb B, and has redescribed
FIGURE 1, asci and ascospores of Mycosphaerella denigrans ex type: left
side, redrawn from Tomilin (1979, fig 39). (scale = 10um)
136
and illustrated this species. The dimensions given by Tomilin for M.
denigrans are as follows: ascomata 80-100um diam.; asci 30-34 x
10-11pm; ascospores 10-12 x 2.5-3um. | likewise have examined the
holotype of Sphaerella denigrans and found asci and ascospores
which agree with Kirschstein’s description and match the dimensions
and_ illustrations provided by Tomilin for this species. My
measurements for the asci and ascospores were 30-37 x 9-11um
and 12-14 x 3-3.5um_ respectively. The ascomata were
subepidermal, medium brown in colour and 99-110”um wide. The
beak was 30-40jim across and the ostiole ca 10um. M. denigrans
is very similar to the earlier published M. pyrenaica (Speg.) Arx. The
dimensions and morphology of the ascospores of the two species
are very close.
Mycosphaerella densa (E. Rostrup) Lind, Rep. Sci. Results Norw.
Exped. Novaya Zemlya 1921, 19: 12. 1924.
S. densa E. Rostrup, Bot. Tidsskr. 14(3): 225. 1885.
This species was redescribed and illustrated by Corlett (1988, p.
65). The description provided was based in part on observations of
the type collection, ex herb. C.
Mycosphaerella hambergii (Romell & Saccardo) Petrak, Acta Horti
Gothob. 17: 137. 1947.
Sphaerella confinis P.A. Karsten, Ofvers Forh. Kongl. Svenska
Vetensk.-Akad. 1872(2): 106. 1872 subspecies hambergii L.
Romell & Saccardo in Saccardo, Syll. Fung. 11: 296. 1895
Romell and Saccardo (op. cit., 1895, p. 296) described Sphaereila
confinis var. hambergii on Arabis gerardi Bess. from Sweden without
illustrations as follows: "Peritheciis dense gregariis, subsphaeroideis,
100 yp diam., epidermide arcte velatis erumpentibusque, nigris,
subastomis; ascis subclavatis sessilibus, 25-50 x 7-10; sporidiis
oblongo-fusoideis, obscure uniseptatis, plerumque 4-guttatis, rectis
curvulisve, 12-15 x3-4, ex hyalino chlorinis". The dimensions given
for subspecies hambergii are slightly less than those given by
Karsten (op. cit., 1872) for Sphaerella confinis Karsten (= M. tassiana
(de Notaris) Johans.). Tomilin (1979) did not discuss M. hambergii.
| have not seen any material of this species and am unable to
comment on its taxonomic status.
137
Mycosphaerella isatidis Kalymbetov, Trudy Inst. Bot.7: 326. 1959.
M. isatidis was described by Kalymbetov (op. cit., 1959, p. 326)
on /satis tinctoria L. from Kazakhstan. Kalymbetov provided the
following description without illustrations: “Perithecia globosa, nigra,
primo, immersa, dein emergentia. Asci cylindrici, basi fasciculati,
hyalini, aparaphysati, 12,3-21,5 x 34,4-51,6 yu. Sporae hyalinae,
bicellulares, cylindricae, 8,6-12,3 x 4,3-6,4 yu". Tomilin (1979, p. 291)
provided a brief description of Kalymbetov’s species without
illustrations: ascomata 70-100um; asci 34-52 x 12-21~7m; ascospores
10-13 x 4-5(6)um; host /satis tinctoria L. This species is known to
me only from the literature. | have not seen type or any other
material of M. isatidis and am unable to comment on its taxonomic
status.
Sphaerella napicola Fautrey, Rev. Mycol. 12(46):64. April 1890.
S. napicola was described by Fautrey (op. cit, 1890, p. 64) on
dead stems of Brassica napus L. [var. oleifera?]. Fautrey provided
the following description without illustrations: “Taches grandes
indéterminées noires; Péritheces nombreux, pressés sur la tache et
disséminés au dehors, arrondis, noirs; Theques ventrues; Spores
hyalines, uniseptées, a 4 gouttes, a loge supérieure plus longue et
plus large, 18, 28 x 4, 6 [18.28 x 4.6]".
Tomilin (1979, p. 77) listed S. napicola as a synonym of
Mycosphaerella cruciferarum (Fries) Lindau. Certainly, the
ascospore measurements given by Fautrey are within the range of
those of M. cruciferarum. | have not seen the type or any other
collections of this species.
Mycosphaerella pachyasca (E. Rostrup) Vestergren, Bih. Kong.
Svenska Veten.-Akad. Handl. 26(3, 12): 7. 1900.
Sphaerella pachyasca E. Rostrup, Meddl. Groenland 3(2): 552.
1888. FIG. 2
Rostrup (op. cit., p. 552-3) stated that Sphaerella pachyasca
Rostrup was common and widespread, occurring on dicotyledonous
138
members of numerous plant families in Greenland, including Draba
and Arabis, members of the Brassicaceae. He considered it to be
the morphological equivalent of Sphaerella tassiana de Notaris, the
latter being restricted to monocot hosts by Rostrup. Further,
Rostrup stated that these two species or species complexes were
probably converging and in many cases could only be distinguished
on the basis of whether the host was a dicot or a monocot [English
translation of Rostrup’s discussion provided by Dr. H. Knudsen,
Keeper of Mycology at the Botanical Museum (C)]. Rostrup
described S. pachyasca without illustrations as follows: “Perithecia
Sparsa v. gregaria, foliicola v. cladogena. Asci crasse ovoideo-
oblongati, inaequilaterales, long. 40-50 p/, crass. 15-24 py, vulgo apice
tunicati. Sporae conglobatae, conoideo- v. ovoideo-oblongatae, long.
16-20 py, crass. 5-6 yp.
Von Arx (1949, p. 40) considered this species to be a synonym
of Mycosphaerella tassiana (de Notaris) Johanson. | have
FIGURE 2, asci and ascospores of Mycosphaerella pachyasca from one of
several collections labelled "Typus", Disco |., 20-7-84[1884]. (scale = 10um)
139
examined collections of S. pachyasca cited by Rostrup (op. cit., p.
552) ex Herb. C and all labelled as “TYPUS". | found asci and
ascospores representative of S. pachyasca in only a few of these;
the asci and ascospores measured 56-60 x 17-17.5um and 15-20 x
5-6.2um respectively. These collections are morphologically
indistinguishable from material | consider to be representative of M.
tassiana (Corlett, 1988). | agree with the use of M. tassiana sensu
von Arx as a single broad unspecialized species or species complex
perhaps, widespread on many monocot and dicot plant families. If
M. pachyasca were to be maintained as a distinct species, a
lectotype would have to be selected from among the collections
listed by Rostrup with his description of S. pachyasca.
Collections examined and yielding ascospores: On Plantago borealis,
Itivnek, W. & H., 13-7-84. On Diapensia lapponica, Disco |., W. & H.,
20-7-84. On Draba hirta, Upernivik, 8-84.
Mycosphaerella pashkiensis Petrak, Sydowia 16(1-6): 340. 1962 [May
1963]. FIG. 3
M. pashkiensis was described by Petrak (op. cit., p. 340) on dead
leaves of Draba species from Afghanistan. Petrak provided the
following description without illustrations: "Perithecia amphigena sed
plerumque hypophyila, per totam folii sSuperficiem irregulariter
dispersa, solitaria, raro bina complurave subaggregata,
subepidermalia, depresso-globosa vei late ellipsoidea, 70-100 p
diam., raro etiam paulo majora, omnino clausa, nec ostiolata; pariete
membranaceo, ca. 10 pt crasso, contextu pseudoparenchymatico e
cellulis irregulariter angulosis vel fere globosis, vix Ccompressis,
obscure atro-brunneis, 3,5-6 pj diam. metientibus Composito; asci sat
numerosi, rosulati, cylindracei, antice late rotundati, postice plus
minusve sed plerumque parvum saccati, subsessiles vel brevissime
et crassiuscule noduloso-stipitati, 8-spori, 42-55 x 9-10,5 ul; sporae
di-vel indistincte tristichae, clavato-cylindraceae vel bacillari-clavatae,
utrinque obtusae, antice vix vel parum., postice plerumque distincte
et paulatim attenuatae, rectae, raro inaequilaterae, circa medium
septate nec constrictae, guttulis plerumque duabus in quaque cellula
praeditae, hyalinae, 10-13 y, raro usque ad 14,5 py longae, 2,5-3,5
pi latae; paraphysoides paucae, indistincte fibrosae, mox omnino
mucosae". Mycosphaerella pashkiensis Petrak was accepted by
Tomilin (1979, p. 76) who published a brief description without
140
illustrations: ascomata 70-100um; asci 42-55 x 9-10.541m; ascospores
(10)12-14 x 2.5-3(3.5)um; host Draba.
| have examined the type collection of M. pashkiensis ex herb. W.
My dimensions of the asci (specifically ascus width) and ascospores
from the type collection exceed the measurements given by Petrak
for M. pashkiensis. My description of the type material is as follows:
ascomata subepidermal, medium brown, 78-108um diam., ostiole,
pale brown, ca 10um diam. Asci bitunicate, thick-walled, saccate
when young, clavate to somewhat cylindrical, widest above or below,
8-spored, 42.5-51 x 12.5-14um [Petrak: 42-55 x 9-10.5].
Ascospores hyaline, mostly cylindrical-clavate, straight or curved, 15-
18.5 x 4-5.25um [Petrak: 10-13(14.5) x 2.5-3.5], septate at or near
the middle, not or hardly constricted at the septum, upper cell wider
and ascospores distinctly tapering to the base, ends rounded but
not broadly so, bi- to triseriate in the ascus. | believe that both
Petrak and | observed the same fungus but | cannot account for the
discrepancy in our dimensions of the asci and ascospores. My
measurements approach those of a smail or immature
Mycosphaerella tassiana.
FIGURE 3, ascoma, asci and ascospores of Mycosphaerella pashkiensis ex
type. (scales = 100um & 10um respectively)
141
Mycosphaerella pyrenaica (Speg.) Arx, Sydowia 3:64. 1959.
Sphaerella pyrenaica Speg., Rev. Mycol. 4(14): 78. 1882.
FIG. 4
Spegazzini (op. cit. 1882, p. 78) described S. pyrenaica from dead
tissue of Peterocallis [Draba]] pyrenaica from France. He provided
the following description without illustrations: "Perithecia amphigena,
globoso-lenticularia (90-100), innato-erumpentia, atra,
submembranacea, contextu§ parenchymatico, fuligineo; asci
cylindraceo-clavati, breve stipitati (40-45 x 8), apice crassiuscule
tunicati, octospori, aparaphysati; sporidia disticha, cylindraceo-
elliptica, utrinque obtusiuscule rotundata (15 x 3), medio 1-
pseudoseptata, non constricta, hyalina". The dimensions given by
von Arx (op. cit., p. 64) generally agreed with those of Spegazzini:
“Pseudothecien ... 90-130 pu... Asci ... 40-50 x 9-11 yw... Sporen ...14-
19 x 3-4 yp ..". M. pyrenaica was accepted by Tomilin (1979, p. 77)
who listed Cardamine bellidifolia L., Cochlearia officinalis L., Parrya
nudicaulis (L.) Regel and Petrocallis [Draba] pyrenaica as hosts.
| have examined the type
collection of S. pyrenaica
Speg., ex herb. LP and
concur with the description
and dimensions provided by
Spegazzini and by von Arm.
My observations of type
material are as_ follows:
ascomata subepidermal, 85-
100um diam, medium brown;
asci__cylindrical-clavate, 8-
spored, 40-48 x 9.5-12um;
ascospores hyaline, cylindrical-
ellipsoidal, straight
FIGURE 4, ascus and ascospores
of Mycosphaerella pyrenaica ex
type (scale = 10um)
or curved, 14.5-18.5 x 3.5-4.5um, septate at or just below the middle,
not or hardly constricted at the septum, upper cell sometimes
slightly wider than lower cell, ends rounded, bi- to triseriate in the
asSCuUS.
142
Mycosphaerella tassiana (de Notaris) Johanson, Ofvers. Férh. Kongl.
Svenska Vetensk.-Akad. 41(9): 167. 1884.
Sphaerella tassiana de Notaris, Sferiacei Italici 1(1): 87. 1863.
Anamorph: Cladosporium herbarum (Pers.) Link: S.F. Gray,
Nat. Arr. Brit. Pl. 1:556. 1821.
A very common and distinctive species found on numerous host
plants in different monocot and dicot families. Lind (1934, p. 60)
listed 110 species, a few non-angiosperms but mostly monocots and
dicots, as hosts of M. tassiana. This species has been described
and illustrated by von Arx (1949, pp. 40-59), M.E. Barr (1959, p. 23-
27; 1972, p. 603), Sivanesan (1984, p. 225) and Corlett (1988, p.
67-72). An extensive list of tassiana synonyms, including M. allicina
(Fr.: Fr.) Vestergren and M. pachyasca (E. Rostrup) Vestergren (see
above) and a lengthy host list were provided by von Arx (op. Cit.)
while de Vries (1952) listed the synonyms of the anamorph. Tomilin
(1979, p. 270) also considered tassiana and allicina to be synonyms
but unlike von Arx, listed M. tassiana as a synonym of the earlier
published M. allicina (basionym: Sphaeria allicina Fr.: Fr., Kongl. Vet.-
Akad. Handl. 38:247. 1817). M.E. Barr (1972, p. 600) maintained
the morphologically similar M. allicina as distinct, considering it to be
specialized and host restricted, parasitic on species of Allium in
contrast to the wide unspecialized host range of M. tassiana.
Mycosphaereila _vesicariae-arcticae (Hennings) Tomilin,
Opredelitle’gribov roda Mycosphaerella Johans., p. 78.
1979.
Sphaerella vesicariae-arcticae P. Hennings in Allescher &
Hennings, Biblioth. Bot. 42: 45. 1897.
This species was described by Hennings (op. cit. 1897, p. 45) on
leaves and stems of Vesicaria arctica R. Br. [this authority differs
from those of the various homonyms of Vesicaria arctica given in
Index Kewensis] var. /ejocarpa Trautw. from Denmark. Hennings
provided the following description without illustrations: “Peritheciis
Caulicolis et amphigenis .sparsis gregariisve, subglobosis atris; ascis
oblongis subclavatis vel subfusoideis 60-75 x 17-21 tu, apice obtusis,
8-sporis; sporidiis subdistichis vel conglobatis, clavatis utrinque
obtusis, medio 1 septatis haud constrictis, rectis, hyalinis 20-25 x 7-
8 yu". Tomilin (op. cit., 1979, p.78) transferred Hennings’s species to
Mycosphaerella and provided a description without illustrations:
143
ascomata 90-120um; asci 60-65 x 20-24y1m; ascospores 17-19 x
(5.5)6-7ym. Tomilin listed Vesicaria and Parrya nudicaulis (L.)
Regel. as hosts. | have not seen the type or any other specimens
of vesicariae-arcticae although the descriptions provided by both
Hennings and Tomilin suggest a small spored Mycosphaerella
tassiana.
144
Dimensions of ascomata, asci and ascospores of
species of Mycosphaerella on Brassicaceae taken
from original authors or from Corlett© (1988)
Name Ascomata diameter ASsci Ascospores
alysii 50-70 24-30x9-10 9-10x3-4
brassicicola® 90-145 30-47(55)x12-16 16-25x3-4
carniolica 30-42x14-18 15-19x4-5
confinis” 150 50-65x11-13 16-24x3-5
cruciferarum® 84-125 34-55x9-16 13-15x3.5-5
denigrans 80-100 30-37x9-11 12-14x3-3.5
densa® 85-140 36-54x9-11 14-24x3-4.5
hambergii 100 25-50x7-10 12-15x3-4
isatidis 34-51x12-21 8.5-12x4-6
napicola”™” 18.28x4.6
pachyasca" 40-60x15-24 16-20x5-6.5
pashkiensis© 78-108 42-51x(9)12-14 [10-13x2-3.5]
15-18x4-5
pyrenaica 85-100 40-48(50)x9-12 14-19x3-4.5
tassiana® 75-155 45-88x15-29(32) 17-31x4.5-9.5
vesicariae 90-120 60-75x17-24 17-25x6-8
-arcticae
" = tassiana
™“ = cruciferarum
145
Host Index
Species of Mycosphaerella on genera of Brassicaceae
Alyssum
alyssi™
Arabis
hambergii
tassiana
Brassica
brassicicola
cruciferarum
napicola
(= cruciferarum)
Braya
confinis™
(= tassiana)
densa
tassiana
Cardamine
densa
pachyasca
(= tassiana)
pyrenaica
tassiana
Cheiranthus
cruciferarum
Cochlearia
densa
pyrenaica
tassiana
Draba
carniolica
confinis
(= tassiana)
cruciferarum
densa
pachyasca
(= tassiana)
pashkiensis™
tassiana
Erysimum
cruciferarum
denigrans™
Eutrema
cruciferarum
tassiana
isatis
isatidis™
Lesquerella
tassiana
Parrya
pyrenaica
tassiana
vesicariae-arcticae
Petrocallis (=Draba)
pyrenaica™
Vesicaria
vesicariae-arcticae™
™ = Type Host
146
Acknowledgements
We would like to thank Dr. M.E. Barr Bigelow, Sidney, British
Columbia and Dr. S.J. Hughes, C.L.B.R.R., Ottawa, for critically
reviewing the manuscript. | am grateful to the directors and
curators of the following herbaria for the loan of specimens in their
keeping: B, C, LP, UPS & W. | wish to thank Dr. H. Knudsen,
Keeper of Mycology at the Botanical Museum (C) for kindly providing
an English translation of Rostrup’s discussion following the
description of Sphaerella pachyasca Rostrup.
References
Ar, J.A. von. 1949. Beitrage zur Kenntnis der Gattung Mycosphaerella.
Sydowia 3: 28-100.
Barr, M.E. 1959. Northern Pyrenomycetes I. Canadian Eastern Arctic.
Contr. Inst. Bot. Univ. Montréal 73:1-101.
Barr, M.E. 1972. Preliminary studies on the Dothideales in temperate North
America. Contr. Univ. Mich. Herb. 9(8): 523-638.
Corlett, M. 1988. Taxonomic studies in the genus Mycosphaerella. Some
species of Mycosphaerella on Brassicaceae in Canada. Mycotaxon
31(1): 59-78.
De Vries, G.A. 1952 Contribution to the knowledge of the genus
Cladosporium Link ex Fr., 121 pp., Diss. Univ. Utrecht, Baarn.
Lind, J. 1934. Studies on the geographical distribution of arctic circumpolar
micromycetes. Kongel Danske Videnskabernes Selskab. Biol.
Meddel. XI, 2 pp. 54-72.
Punithalingam, E. & P. Holliday. 1975. CMI Descript. Path. Fungi & Bacteria
No. 468. Mycosphaerella brassicicola. CAB Intern. Mycol. Inst., Kew,
Surrey.
Sivanesan, A. 1984. The bitunicate Ascomycetes and their anamorphs. J.
Cramer, Vaduz, pp. 1-701.
Tomilin, B.A. 1979. Opredelitel’gribov roda Mycosphaerella Jonans., Nauka,
Leningrad, pp. 319.
MY COTAXON
Volume XLVI, pp. 147-155 April-June 1993
DEVELOPMENT OF CONIDIOMATA IN THE PHYLLOSTICTA STATE OF
GUIGNARDIA MANGIFERAE ROY AND OBSERVATIONS ON THE
FINE STRUCTURE OF THE CONIDIUM
by
J. Muthumary, J.A. Jayachandra and M. Bhagavathy Preetha
Centre for Advanced Studies in Botany
University of Madras, Madras-600 025, INDIA
SUMMARY
The development of conidiomata in the Phyllosticta state of Guignardia
mangiferae Roy on the host and in culture is described. Fine structural details of the
sheath around the conidium is provided.
INTRODUCTION
The developmental stages of the conidiomata in Coelomycetes have only been
studied in detail in a few members of this group (Punithalingam 1966;1981;1982; Maas
et al., 1979; Muthumary and Vanaja 1986). In this investigation developmental stages of
the conidiomata in the Phyllosticta state of Guignardia mangiferae Roy were investigated.
This forms part of a programme of work on developmental morphology and taxonomy of
coelomycetes which is in progress in this department. An attempt was also made to
provide information regarding the fine structure of the conidium.
MATERIALS AND METHODS
The material used in this study was collected in Vandaloor, near Madras, growing
on leaves of Mangifera indica in September, 1991. The fungus was isolated in pure
culture on oatmeal agar. The specimen was processed for TEM studies with a view to
obtaining information on the ultrastructural details of the conidium and its appendage in
relation to the wall layers. Portions of agar with conidiomata were fixed in primary
fixative ie. 5.0% Glutaraldehyde (W/V) in 0.1 M phosphate buffer (pH=7.5) for 30 min.
at room temperature and for 1 hr,. at 4°C. Similarly pieces of leaf with conidiomata were
also processed in the same manner. Three washes were given in the buffer solution at 4°C
to remove traces of the GA, before fixing in 1.0% O,O, for 1 hr at 4°C. After washing
several times in phosphate buffer the specimen was dehydrated through a graded series
of Acetone. Subsequently the specimen was embedded in Araldite (Araldite My
148
753=10ml, DDSA=10 ml, BDMA = 0.4 ml, N.Dibutylphthalate =0.5m1) and polymerized
at 60°C for 36 hrs. Gold colour sections were cut with an ultracut ultramicrotome.
Semithin sections of about 0.1 » thickness were selected to follow the different stages of
development of the conidiomata. The semithin sections were stained with aqueous
Toluidine blue for light microscopic observations. Ultrathin sections were collected on
copper grids and stained with Uranyl acetate followed by lead citrate (Reynolds, 1963).
Sections were observed with a CM 10 Philips Electron Microscope at 40 Kv.
RESULTS
Formation of conidiomatal primordium. The earliest stage of pycnidial
development seen in sectioned material from the culture consists of a small,
pseudoparenchymatous mass of tissue, the pycnidial primordium (Fig. 1). The pycnidial
primordium is almost spherical, consisting of a compact cluster of brown, thick -walled
cells. During further development, the primordium considerably increases in size by
continued transverse and longitudinal divisions of the cells.
Formation of the pycnidial cavity. As a prerequisite for the formation of the
pycnidial cavity, the cells at the periphery and centre of the primordium were arranged
as follows. The cells along the outer layers of the primordium became arranged
tangentially which subsequently formed the conidiomatal wall (Figs. 2-4). The inner cells
of the primordium were thin-walled and hyaline whereas the peripheral cells were brown
and thick - walled. There may be more than one method involved in the formation of the
central cavity. The presence of ruptured and disorganised cells in the cavity indicates that
the cavity might have formed by the rupture of cells at the centre of the primordium
(Figs. 5-6). The occurrence of hyphal fragments and a gelatinous material within the
Cavity suggest that the cells are later dissolved to form the cavity (Fig.3). From the many
sections examined, both these conditions were observed as being present in the fungus.
The tangentially arranged outer layers of cells form the pycnidial wall by repeated
division, thereby increasing the dimension of the developing conidioma. Simultaneously,
the outer wall layers become carbonaceous and at maturity the wall consists of 5-6
tangentically arranged layers, the cells of the inner layers being hyaline and thin-walled
(Figs. 10-11).
Formation of sporogenous tissue. In conidiomata formed in culture, the
formation of conidia from the conidiogenous cells resembles that of Ascochyta spp..
(Punithalingam, 1979) and Macrophomina phaseolina (Tassi) Goid. (Punithalingam, 1982).
After the cavity is formed by dissolution and disintegration of central cells of the
primordium, the cells lining the cavity commence conidial production by a budding like
process. Eventually these conidiogenous cells, along with conidia become detached from
the wall of the conidioma (Figs. 5- 8), and remain floating still intact in the conidiomatal
cavity (Figs 7-8). The conidiogenous cells which were produced initially inside the cavity
are referred to as "temporary conidiogenous cells" by Punithalingam (1979). These
temporary conidiogenous cells very occasionally bear an annellation and were normally
blastic. Subsequently the cells lining the cavity elongate to form a palisade of permanent
conidiogenous cells (Punithalingam, 1979) which then start producing conidia (Fig. 9).
149
The presence of permanent conidiogenous cells were observed even when the cavity filled
up with dissolved and disintegrated primordial cells (Fig. 9). Consequently the conidioma
becomes considerably larger in size by proliferation of the outer layers of the
pseudoparenchyma while the inner cells become arranged tangentially into several layers
(Figs. 10-12).
Formation of ostiole. When the conidioma becomes fully mature, a papilla-like
projection appears at the apex (Figs. 16-18). In vertical section, the papilla shows
thick-walled, dark cells towards the outer side and thin walled, hyaline, elongated cells
towards the inner side (Figs. 13-15). During dehiscence of the conidioma, these inner
hyaline layers in the papilla dissolve to form the ostiole and the outer, thick walled cells
of the papilla remain intact and form the papillate structure of the conidioma. The papilla
may also exert pressure to rupture the epidermis of the host for releasing the conidia
outside.
Fine structure of the conidia. Light microscopic studies by Punithalingam
(1982) showed that the conidium is enveloped by a thick sheath which remains intact, its
outer most layer gradually gelatinizing with the ageing of the conidium, and our studies
using TEM confirmed the presence of this thick sheath around young conidia. Our
observations showed the presence of the sheath around the conidiogenous cells and the
growing conidium (Fig. 20). Transverse sections of a young conidium also revealed the
presence of the sheath (Figs, 22-23 & 25), but in TEM micrographs of old conidia the
sheath around the conidia was absent (Figs. 19, 21 & 24). It is therefore probable that
during the later stage of development, the sheath around the conidium gradually
gelatinizes. In observations with the light microscope, conidia within young pycnidia were
found with the sheath surrounding them (Fig. 14). The mature conidium is multinucleate
(Punithalingam and Woodhams, 1982) and in this study, at least three nuclei were
observed (Fig. 21). TEM studies showed that the sheath surrounding the conidium
disappears with ageing (Figs. 19, 21).
DISCUSSION
The orgin and structure of the apical conidial appendage of Phyllosticta spp. had
already been fully described by Punithalingam and Woodhams (1982). The cultural
characters of the Phyllosticta state of Guignardia mangiferae Roy along with the
connection between the ascogenous state has also been thoroughly investigated
(Punithalingam 1974). This work deals with the different stages of development of the
conidiomata within the natural host and also in artificial culture media. It was observed
that the Phyllosticta sp. investigated here produces temporary and permanent
conidiogenous cells similar to those of Ascochyta spp. (Punithalingam 1979) and
Macrophomina phaseolina (Punithalingam, 1982). Normally the temporary conidiogenous
cells produce a single conidium after which they get sloughed off from the wall layers.
In the present fungus, the temporary conidiogenous cells occasionally show one
annellation indicating that it is capable of producing more than one conidium. Probably
after producing one or two conidia, the temporary conidiogenous celis may become
detached from the wall layers.
150
Another interesting finding is the presence of the thick sheath around the
conidium. This sheath was also observed covering the conidiogenous cell and the
conidium initial. The presence of the sheath around the conidium confirms observations
made by light microscopy. Both transverse and longitudinal sections of the old conidia
failed to reveal a sheath surrounding the conidia suggesting that the sheath gradually
gelatinizes and disappears as the conidium becomes old.
The dual conidiation process was found as a general feature of Ascochyta species
in culture (Punithalingam 1979) and also in Macrophomina phaseolina (Punithalingam
1982). Other Coelomycete species studied by the authers, namely, Botryodiplodia
theobromae Pat., Ciliochorella mangiferae Syd., Coleophoma cylindrospora (Desm.)
Hohn., Pestalotiopsis palmarum (Cooke) Stey. and Urohendersonia pongamia Naj Raj
& Ponnappa also show this type of dual conidiation process in culture. In the present
investigation the Phyllosticta species also was found to produce temporary and permanent
conidiogenous cells. But Robillarda depazeoides (Welw. et Curr.) Sacc. was found to
produce only one type of conidiogenous cells during conidiation in culture. Therefore the
present study reveals that necessary data collected based an the developmental
morphology of conidiomata for many more Coelomycetes will be useful for segregating
genera into differant groups. In addition, the ultrastructural details of the conidia given
in the present investigation have provided useful’ information regarding the biological
significance of the mucilaginous sheath around the conidium.
REFERENCES
Maas, J.L., Pollack, F.G. and Uecker, F.A. 1979. Morphology and development of
Pilidiella quercicola. Mycologia 71: 92-102.
Muthumary, J. and Vanaja,R. 1986. Development of conidiomata in Coniella fragariae.
Trans. Br. Mycol. Soc. 87(1): 109-114.
Punithalingam, E. 1966. Development of the pycnidium in Septoria. Trans. Br. Mycol.
Soc. 49: 19-25,
aona----- , 1974. Studies on Sphaeropsidales in culture II. Mycol. Pap. 136: 1-63.
wennnnnee , 1979. Graminicolous Ascochyta Species. Mycol. Pap. 142: 1-214.
wonnenn-- , 1981. Conidiation and appendage formation in Tiarosporella paludosa (Sacc. &
Fiori) Hohnel. Nova Hedwigia 34: 539-567.
--------- , 1982. Conidiation and Appendage formation in Macrophomina phaseolina (Tassi)
Goid. Nova Hedwigia 36: 249-290.
w-------- and Woodhams, J.E. 1982. The conidial appendage in Phyllosticta spp. Nova
Hedwigia 36: 151-198.
Reynolds, E.S. 1963. The use of lead citrate at high pH as an electron opaque stain in
electron microscopy. Journal of cell Biology. 17: 208-212.
151
Fig.1. Transverse section of pycnidial primordium x 300 Figs.2-4. Transverse sections of
young primordia showing the formation of central cavity. x 300 Figs.5-6. Transverse
sections of young conidiomata showing the central cavities with the disentegrated hyphal
fragments. x 300
152
oS Se
Figs.7-8. Central cavities showing the sloughed off temporary conidiogenous cells with
the conidia. x 528. Fig.9. Section of conidioma showing the formation of permanent
conidiogenous cells. Note the presence of detached, temporary conidiogenous cells. x 2646
Figs.10-12. Vertical sections of mature conidioma. Fig.10 x 264; 11 x 132; 12 x 88.
Figs.13-15. Vertical sections of conidicmata on leaf showing the papillae piercing the
epidermis of the host 13 x 350, 14 x 350, 15 x 183. Figs. 16-18. Vertical sections of the
conidiomata on leaf showing the papillae. 16 x 350, 17-18 x 183.
Figs. 19-25. Transmission electron micrographs of the Phyllosticta state. Bar = 3um.
Fig. 19. Section of mature conidium. Note the absence of the sheath surrounding the
conidium. Fig. 20. Section of a young conidiogenous cell and conidium. Note the
presence of the sheath around the conidiogenous cell and the conidium initial. Fig.
21. Section of a mature conidium showing three nuclei. Note the absence of a sheath
around the conidium. Fig. 22. Section of a you ng conidium showing the sheath
around the wall. Fig. 23. Section of more or less a mature conidium showing
remnants of the sheath around the wall.
15D
Figs. 24-25. Bar=3ym. Fig. 24. Section of a mature conidium showing remnants of
the sheath around the conidium. Fig. 25. Section of a young conidium showing the
sheath.
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MY COTAXON
Volume XLVI, pp. 157-159 April-June 1993
HYDNANGIUM PILA Pat., AN OLDER NAME FOR MARTELLIA
MEDITERRANEA Moreno, Galan & Montecchi
G. MORENO & R. GALAN
Department of Plant Biology, Alcala de Henares
University, 28871 Madrid, Spain
In a recent publication (MORENO, GALAN & MONTECCHI,
1991), a new species belonging to the hypogeous fungi was
described, Martellia mediterranea (Russulales), having been found
in Spain, growing with Quercus suber and Cistus ladanifer. It was
compared with Zelleromyces stephensii (Berk.) A. Smith and Martellia
mistiformis Matt.
It. was very surprising for us to note that at almost
the same time in a local bulletin (VIDAL, 1991), a fungus
"apparently" cospecific with the refered species was
described in detail, identified as Hydnangium pila
(PATOUILLARD, 1910) and with the proposal of the new
combination: Martellia pila (Pat.) Vidal for it, as well as a
list of possible different synonyms.
In order to clarify this hypothetical taxonomical
controversy, we decided to study a duplicate of Vidal's
material (now in the private herbarium of J. M. Vidal, ref.
910523-3 and 901226-6), which was kindly sent to’ us by Mr.
Vidal, as well as the type of Hydnangium pila Pat., which was
received from the Farlow Herbarium (Harvard University,
Cambridge) -Fig.1-, thanks to the kindness of Dr. D. H.
Pfister.
After carefully comparing those specimens with Martellia
mediterranea, we must conclude that our suspicions were
certain and all of them represent the same _ species,
correctly named Martellia pila (Pat.) Vidal.
At the same time we consider convenient to point out
that the type of MHydnangium pila, consisting of three
basidiocarps (two immature and the other one well
preserved) shows a wide peridium -reaching 200-300 pm in
diam. - with no sphaerocysts and having a remarkable
gelification (Figs. 5 & 6). On the other hand, basidia are
tetrasporic -exceptionally bisporic- measuring 28-40 x 14-
17 pm, and cystidia were not observed, probably due to the
age of the fructification, according to Vidal's suggestion.
ins sspite, sof yithis™ the strongly amiloid and _ spiny
basidiospores, 9-11 % 9-10 wpm, are good diagnostic
characters (Figs. 2-4).
158
ACKNOWLEDGEMENTS
We wish to express our gratitude to Mr. Josep M. Vidal
and Dr. Donald H. Pfister for loan of specimens and to Dr.
Richard P. Korf for critical review of the manuscript, as
well to Dr. F. Esteve for help in preparing the English
text.
LITERATURE CITED
MORENO G., GALAN, R. & MONTECCHI, A. (1991). Hypogeous
fungi from Peninsular Spain. II. Mycotaxon 42:201-238.
PATOUILLARD, N. (1910).Note sur trois espéces
d'Hydnangium de la flore du Jura. Bull. Soc. Mycol. France
26:199-204.
VIDAL, J. M. (1991). Contribucién al conocimiento de la
flora micolégica del Baix Emporda y zonas'7 limitrofes
(Catalunya). IV. Hongos hipogeos (Zygomycotina, Ascomycotina
y Basidiomycotina). Butll. Soc. Catalana Micol. 14-15:143-
194.
Figs. I-6. Hydnangium pila Pat., type: Fig. 1. Original label
handwritten by Patouillard. Figs. 2-4. Basidiospores.
Fig.5. Cross-section of a sporocarp under light microscope.
Fig.6. Texture of the peridium.
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MY COTAXON
Volume XLVI, pp. 161-176 April-June 1993
COMPUTER CODING OF STRAIN FEATURES
OF THE GENUS RHIZOPUS
SHUNG-CHANG JONG! and CANDACE MCMANUS?
‘American Type Culture Collection, 12031 Parklawn Drive, Rockville, MD
20852 USA and 7Microbial Systematics Section, National Institute of Dental
Research, National Institutes of Health, Bethesda, MD 20892 USA
ABSTRACT
Members of the fungal genus Rhizopus play important roles in human, animal,
and plant disease and in the fermentation industry. Identification of Rhizopus
species is based primarily on the morphology of sporangiosporogenous structures,
Zygospores, and asexual reproductive structures. The standardized coding system,
the RKC Code, used for computer storage and analysis of microbial strain data
was expanded to include features specific to the identification of Rhizopus species.
BACKGROUND AND DISCUSSION
Members of the genus Rhizopus occur as saprophytes on plant debris
in soil. Many are known to cause soft rots or spoilage of vegetables
and fruits. Some are etiologic agents of mucormycosis in humans and
animals. Species of Rhizopus play important roles in the
fermentation industry, especially in the production of enzymes (e.g.,
glucoamylase and lipase), organic acids, alcohol, and fermented foods
such as bongkrek, lao-chao, tauco, and tempeh (Inui et al., 1965;
Alexopoulos and Mims, 1979).
The genus Rhizopus belongs to the family Mucoraceae under the
order Mucorales. The genus was established by Ehrenberg (1820)
and its history first reviewed by Zimmermann (1871). Rhizopus
taxonomy has been the subject of many well known reports including
those of Fischer (1892), Lendner (1908), Hanzawa (1915), Zycha
(1935), Zycha etal. (1969), Naumov (1939), Inui et al. (1965),
162
Dabinett and Wellman (1973), Schipper (1984), Schipper and Stalpers
(1984), and Liou et al. (1990).
The taxonomy of Mucorales has been based primarily on
morphological characteristics of sporangiosporogenous structures and
zygospores. The genus Rhizopus, based on the type species R.
stolonifer (Ehrenberg:Fries) Vuillemin, is characterized by mycelia
bearing stolons and rhizoids and by formation of sporangiophores
arising from the points of rhizoid attachment. Sporangia are
differentiated at the tips of sporangiophores and are spherical
(globose) with columellae and apophyses. Sporangiospores are
produced within the sporangium outside the columellae and are
mostly regular in shape and striated (Hesseltine and Ellis, 1961).
Although the zygospore is the major morphological characteristic
used for species delineation, asexual reproductive structures are most
commonly used in routine identification of Rhizopus cultures.
Since the genus Rhizopus was established, more and more differential
characters were used for species delineation, and, as a consequence,
many species in the genus were described. However, species
differentiation was often, based on minor differences. In a numerical
taxonomic analysis of data previously published by Inui et al. (1965),
Dabinett and Wellman (1973) found that many of the 34 taxa
included in the analysis were very closely related and that few of the
133 physiological and 16 morphological characters contributed
significantly to the classification. More recently, other techniques
have been used to study the taxonomy of the genus Rhizopus,
including scanning electron microscopy of spore ornamentation
(Schipper and Stalpers, 1984), mating experiments (Schipper, 1984;
Schipper and Stalpers, 1984), and DNA homology studies (Ellis, 1985,
1986).
Many previously described Rhizopus species are now considered to be
synonymous, and the number of recognized species has been reduced
considerably. Schipper (1984) and Schipper and Stalpers (1984)
recognized only five species. Only two species were retained in the
R. stolonifer-group -- R. stolonifer and R. sexualis, each with two
varieties. In the R. microsporus-group, two species were also retained
-- R. homothallicus and R. microsporus, the latter with four varieties.
163
The fifth species was R. oryzae. An additional species, R. azygosporus,
was subsequently described by Yang and Jong (1984).
Use of a standardized vocabulary for coding microbial strain data
facilitates computer storage, retrieval, analysis, and exchange of such
data. Data encoded in this manner can be used to build computer
databases for study of taxonomic relationships and construction of
identification keys or probability matrices for computer-aided
identification of new isolates. In this communication, we present a
comprehensive set of features important in the differentiation of
Rhizopus species that have been incorporated into an existing
computer coding system, the RKC Code.
The RKC Code (after the original authors -- Rogosa, Krichevsky, and
Colwell, 1971) is an open-ended, statement-oriented controlled
vocabulary of descriptors of strain characteristics or features. The
RKC Code was developed originally for bacterial strains and later
expanded to include features specific for algae, protozoa, and selected
groups of fungi and to allow coding of characteristics such as
serology, phage typing, properties of nucleic acids, and quantitative
antimicrobial susceptibility (Rogosa et al., 1986). More recently, the
RKC Code was further expanded to include features used to
characterize yeusts (Jong et al., 1988), the saprolegnian fungi (Jong
et al., 1991), and the fungal genera Phytophthora (Jong et al., 1989)
and Pythium (Jong et al., 1992).
The set of characteristics developed for use with Rhizopus species is
presented in the list below. Although some of these features were
already part of the RKC Code, many of the descriptors are unique to
Rhizopus species and were created specifically for this genus. New
features added to the RKC Code for the Rhizopus species are marked
with an "*" in the list. The features include morphological
descriptions of rhizoids, stolons, sporangiophores, sporangia,
columellae, sporangiospores, zygospores, azygospores, suspensors, and
chlamydospores; colony characteristics; and growth temperatures.
The terms used for morphological descriptions are based on the
descriptions given in Hawksworth et al., 1983.
164
Rhizoids
008342:
008417:
*043192:
*043193:
*043194:
*043195:
Stolons
008420:
*043196:
*043197:
*043198:
*043199:
*(043200:
*043201:
* (043202:
*043203:
*043204:
*043205:
*043206:
*043207:
*043208:
*(043209:
*043210:
*043211:
*043212:
*043213:
Hyphae are septate.
Rhizoids (root-like hyphae) are present.
Rhizoid system is complex (highly branched).
Rhizoids are pigmented.
Rhizoids are brown.
Rhizoids are black.
Stolons are present.
Stolons are pigmented.
Stolons are brown.
Stolons are 0.1-1.0 uw wide.
Stolons are 1.1-2.0 u wide.
Stolons are 2.1-3.0 yu wide.
Stolons are 3.1-4.0 uw wide.
Stolons are 4.1-5.0 u wide.
Stolons are 5.1-6.0 u wide.
Stolons are 6.1-7.0 « wide.
Stolons are 7.1-8.0 4 wide.
Stolons are 8.1-9.0 u wide.
Stolons are 9.1-10 u wide.
Stolons are 11-12 yu wide.
Stolons are 13-14 uw wide.
Stolons are 15-16 u wide. -
Stolons are 17-18 yu wide.
Stolons are 19-20 yw wide.
Stolons are > 20 x wide.
Sporangiophores
008539: Sporangia are on sporophores (sporangiophores).
*043214: Sporangiophores occur singly.
*043215: Sporangiophores occur in pairs.
*043216: Sporangiophores occur in groups of 3-4.
*043217: Sporangiophores are straight (erect).
165
008786:
008787:
*043218:
Sporangiophores are branched.
Sporangiophore branching is irregular.
Sporangiophore branching is dichotomous.
*043219:
008792:
008562:
*043220:
*043221:
*043222:
*043223:
*043224:
*043225:
*043226:
*043227:
*043228:
*043229:
*043230:
*043231:
Sporangiophores have swellings.
Swellings are present at branch points of sporangiophore.
Apophyses are present.
Sporangiophores are pigmented.
Sporangiophores are brown.
Sporangiophores are black.
Sporangiophores are < 201 u long.
Sporangiophores are 201-300 uw long.
Sporangiophores are 301-400 wu long.
Sporangiophores are 401-500 yw long.
Sporangiophores are 501-600 uw long.
Sporangiophores are 601-700 yu long.
Sporangiophores are 701-800 wu long.
Sporangiophores are 801-900 uw long.
Sporangiophores are 901-1000 uw long.
Sporangiophores are 1001-1500 wu long.
Sporangiophores are 1501-2000 yu long.
Sporangiophores are 2001-3000 y long.
*043232:
*043233:
*043234:
*043235:
*043236:
*043237:
*043238:
*043239:
*043240:
*043241:
*043242:
*043243:
*043244:
*043245:
*043246:
*043247:
*043248:
*043249:
*043250:
Sporangiophores are > 3000 yu long.
Sporangiophores are 0.1-1.0 uw wide.
Sporangiophores are 1.1-2.0 u wide.
Sporangiophores are 2.1-3.0 u wide.
Sporangiophores are 3.1-4.0 yu wide.
Sporangiophores are 4.1-5.0 wide.
Sporangiophores are 5.1-6.0 4 wide.
Sporangiophores are 6.1-7.0 u wide.
Sporangiophores are 7.1-8.0 u wide.
Sporangiophores are 8.1-9.0 u wide.
Sporangiophores are 9.1-10 uw wide.
Sporangiophores are 11-15 yu wide.
Sporangiophores are 16-20 u wide.
Sporangiophores are 21-25 yu wide.
Sporangiophores are 26-30 u wide.
Sporangiophores are > 30 yu wide.
166
Sporangia
043074:
008811:
008558:
008815:
008563:
008199:
*043251:
008829:
*043252:
*043253:
*043254:
*043255:
008578:
008579:
008580:
008581:
008582:
008583:
008584:
008585:
*(043256:
*(043257:
*043258:
*043259:
*043260:
*043261:
*043262:
*(043263:
*043264:
*043265:
*043266:
*(043267:
*043268:
*(43269:
Sporangia are present.
Sporangia are terminal.
Sporangia are spherical (length to breadth ratio is 1.0-1.05).
Sporangia are prolate spheroidal (length to breadth ratio
is 1.06-1.15).
Sporangial walls are smooth.
Sporangial wall disintegrates concurrently with release of
spores.
Sporangia are powdery.
Sporangia are pigmented.
Sporangia are blue.
Sporangia are brown.
Sporangia are gray.
Sporangia are black.
Sporangia are 1.0-2.0 uw wide.
Sporangia are 2.1-3.0 u wide.
Sporangia are 3.1-4.0 uw wide.
Sporangia are 4.1-5.0 u wide.
Sporangia are 5.1-10 u wide.
Sporangia are 11-15 wu wide.
Sporangia are 16-20 u wide.
Sporangia are 21-30 uw wide.
Sporangia are 31-40 mu wide.
Sporangia are 41-50 mu wide.
Sporangia are 51-60 uw wide.
Sporangia are 61-70 su wide.
Sporangia are 71-80 u wide.
Sporangia are 81-90 u wide.
Sporangia are 91-100 u wide.
Sporangia are 101-150 u wide.
Sporangia are 151-200 yu wide.
Sporangia are 201-250 u wide.
Sporangia are 251-300 su wide.
Sporangia are 301-350 yu wide.
Sporangia are 351-400 mu wide.
Sporangia are > 400 uw wide.
Columellae
008561:
*043270:
*043271:
*043272:
*043273:
*043274:
*043275:
*043276:
*043277:
*043278:
*043279:
*043280:
*043281:
*043282:
*043283:
*043284:
*043285:
*043286:
*043287:
*043288:
*043289:
*043290:
*043291:
*043292:
*043293:
*043294:
*043295:
*043296:
*043297:
*043298:
*043299:
Columellae are formed.
Columellae are spherical (length to breadth ratio
is 1.0-1.05).
Columellae are prolate spheroidal (length to breadth
ratio is 1.06-1.15).
Columellae are broadly ellipsoidal (length to breadth
ratio is 1.16-1.30).
Columellae are ellipsoidal (length to breadth ratio
is 1.31-1.6).
Columellae are conical.
Columellae are applanate (flattened).
167
Columellae are pyriform (pear-shaped, attached at narrow
end).
Columellae are obpyriform (pear-shaped, attached at broad
end).
Columellae are cylindrical.
Columellae are obovoid (egg-shaped, attached at narrow
end).
Columellae have collars.
Columellae are pigmented.
Columellae are gray.
Columellae are brown.
Columellae are 0.1-1.0 u long.
Columellae are 1.1-2.0 uw long.
Columellae are 2.1-3.0 u long.
Columellae are 3.1-4.0 u long.
Columellae are 4.1-5.0 uw long.
Columellae are 5.1-6.0 u long.
Columellae are 6.1-7.0 u long.
Columellae are 7.1-8.0 u long.
Columellae are 8.1-9.0 uw long.
Columellae are 9.1-10 yu long.
Columellae are 11-20 uw long.
Columellae are 21-30 uw long.
Columellae are 31-40 w long.
Columellae are 41-50 wu long.
Columellae are 51-60 yw long.
Columellae are 61-70 yu long.
168
*043300: Columellae are 71-80 yu long.
*043301: Columellae are 81-90 yu long.
*043302: Columellae are 91-100 uw long.
*043303: Columellae are 101-150 yu long.
*043304: Columellae are 151-200 yu long.
*043305: Columellae are > 200 yw long.
*043306: Columellae are 0.1-1.0 u wide.
*043307: Columellae are 1.1-2.0 u wide.
*043308: Columellae are 2.1-3.0 uw wide.
*043309: Columellae are 3.1-4.0 u wide.
*043310: Columellae are 4.1-5.0 u wide.
*043311: Columellae are 5.1-6.0 u wide.
*043312: Columellae are 6.1-7.0 wide.
*043313: Columellae are 7.1-8.0 u wide.
*043314: Columellae are 8.1-9.0 u wide.
*043315: Columellae are 9.1-10 uw wide.
*043316: Columellae are 11-20 wu wide.
*043317: Columellae are 21-30 u wide.
*043318: Columellae are 31-40 u wide.
*043319: Columellae are 41-50 u wide.
*043320: Columellae are 51-60 yu wide.
*043321: Columellae are 61-70 u wide.
*043322: Columellae are 71-80 yu wide.
*043323: Columellae are 81-90 uw wide.
*043324: Columellae are 91-100 mu wide.
*043325: Columellae are 101-150 yu wide.
*043326: Columellae are 151-200 wu wide.
*043327: Columellae are > 200 u wide.
Sporangiospores
008173: Asexual spores (sporangiospores) are produced in sporangia
(spore vesicles).
008592: Sporangiospores occur in masses.
008595: Sporangiospores are spherical (length to breadth ratio
is 1.0-1.05). |
*043328: Sporangiospores are prolate spheroidal (length to breadth
ratio is 1.06-1.15).
169
Sporangiospores are broadly ellipsoidal (length to breadth
ratio is 1.16-1.30).
Sporangiospores are ellipsoidal (length to breadth ratio
is 1.31-1.6).
Sporangiospores are angular.
Sporangiospores are fusiform.
Sporangiospores are irregular in shape.
Surfaces of sporangiospores are striated.
Surfaces of sporangiospores are smooth.
Surfaces of sporangiospores are spiny.
Surfaces of sporangiospores are warty.
*043329:
043047:
*043330:
*043331:
043053:
*043332:
043054:
043056:
043057:
043061:
043068:
043067:
008602:
008603:
008604:
*043333:
*043334:
*043335:
*043336:
*043337:
*043338:
*043339:
*043340:
008597:
008598:
008599:
008600:
008601:
*043341:
*043342:
*043343:
*043344:
*043345:
*043346:
*043347:
*043348:
*043349:
Sporangiospores are pigmented.
Sporangiospores are brown.
Sporangiospores are black.
Sporangiospores are 0.1-1.0 uw wide.
Sporangiospores are 1.1-2.0 u wide.
Sporangiospores are 2.1-3.0 uw wide.
Sporangiospores are 3.1-4.0 u wide.
Sporangiospores are 4.1-5.0 u wide.
Sporangiospores are 5.1-6.0 “4 wide.
Sporangiospores are 6.1-7.0 “ wide.
Sporangiospores are 7.1-8.0 « wide.
Sporangiospores are 8.1-9.0 wu wide.
Sporangiospores are 9.1-10 uw wide.
Sporangiospores are > 10 uw wide.
Sporangiospores are 0.1-1.0 u long.
Sporangiospores are 1.1-2.0 u long.
Sporangiospores are 2.1-3.0 uw long.
Sporangiospores are 3.1-4.0 u long.
Sporangiospores are 4.1-5.0 x long.
Sporangiospores are 5.1-6.0 uw long.
Sporangiospores are 6.1-7.0 u long.
Sporangiospores are 7.1-8.0 u long.
Sporangiospores are 8.1-9.0 uw long.
Sporangiospores are 9.1-10 yu long.
Sporangiospores are 11-12 uw long.
Sporangiospores are 13-14 yu long.
Sporangiospores are 15-16 wu long.
Sporangiospores are 17-18 uw long.
170
*043350:
*043351:
Sporangiospores are 19-20 yu long.
Sporangiospores are > 20 wu long.
Sexual Reproduction
008617:
008618:
008619:
Sexual reproduction occurs.
Strain is homothallic (both mating types on same
mycelium).
Strain is heterothallic (mating types on separate mycelia).
Zygospores
008627:
*043352:
*043353:
008628:
008629:
008630:
*043354:
*043355:
*043356:
*043357:
*043358:
*043359:
*043360:
*043361:
*043362:
*043363:
*043364:
*043365:
*043366:
*043367:
*043368:
*043369:
*043370:
*043371:
Zygospores are produced.
Zygospores are spherical (length to breadth ratio is
1.0-1.05).
Zygospores are prolate spheroidal (length to breadth ratio
is 1.06-1.15).
Zygospores are thick-walled.
Surfaces of zygospores are rough.
Surfaces of zygospores are spiny.
Surfaces of zygospores are warty.
Zygospores are pigmented.
Zygospores are red.
Zygospores are yellow.
Zygospores are brown.
Zygospores are black.
Zygospores are ().1-1.0 « wide.
Zygospores are 1.1-2.0 u wide.
Zygospores are 2.1-3.0 uw wide.
Zygospores are 3.1-4.0 u wide.
Zygospores are 4.1-5.0 u wide.
Zygospores are 5.1-6.0 u wide.
Zygospores are 6.1-7.0 “ wide.
Zygospores are 7.1-8.0 uw wide.
Zygospores are 8.1-9.0 uw wide.
Zygospores are 9.1-10 u wide.
Zygospores are 11-15 yu wide.
Zygospores are 16-20 wu wide.
* (043372:
* 043373:
* (043374:
*043375:
*043376:
*043377:
*043378:
*043379:
*043380:
*043381:
*043382:
*043383:
Zygospores are 21-30 uw wide.
Zygospores are 31-40 yu wide.
Zygospores are 41-50 uw wide.
Zygospores are 51-60 u wide.
Zygospores are 61-70 yu wide.
Zygospores are 71-80 u wide.
Zygospores are 81-90 u wide.
Zygospores are 91-100 yu wide.
Zygospores are 101-150 wu wide.
Zygospores are 151-200 u wide.
Zygospores are 201-250 uw wide.
Zygospores are > 250 mu wide.
Azygospores
*043384: Azygospores (parthenogenetic zygospores) are present.
*043385:
*043386: Azygospores are spherical (length to breadth ratio is
Azygospores are terminal.
1.0-1.05).
i7A
*043387: Azygospores are prolate spheroidal (length to breadth ratio
*043388: Surfaces of azygospores are rough.
*043389: Surfaces of azygospores are spiny.
*043390: Surfaces of azygospores are warty.
*043391:
*043392:
*043393:
*043394:
*043395:
*043396:
*043397:
*043398:
*043399:
*043400:
*043401:
*043402:
*043403:
*043404:
*04340S:
is 1.06-1.15).
Azygospores are pigmented.
Azygospores are brown.
Azygospores are black.
Azygospores are 0.1-1.0 “ wide. .
Azygospores are 1.1-2.0 yu wide.
Azygospores are 2.1-3.0 4 wide.
Azygospores are 3.1-4.0 u wide.
Azygospores are 4.1-5.0 4 wide.
Azygospores are 5.1-6.0 wide.
Azygospores are 6.1-7.0 4 wide.
Azygospores are 7.1-8.0 4 wide.
Azygospores are 8.1-9.0 u wide.
Azygospores are 9.1-10 yu wide.
Azygospores are 11-15 yu wide.
Azygospores are 16-20 u wide.
Lye
*043406: Azygospores are 21-30 u wide.
*043407: Azygospores are 31-40 u wide.
*043408: Azygospores are 41-50 yu wide.
*043409: Azygospores are 51-60 u wide.
*043410: Azygospores are 61-70 yu wide.
*043411: Azygospores are 71-80 su wide.
*043412: Azygospores are 81-90 mu wide.
*043413: Azygospores are 91-100 yu wide.
*043414: Azygospores are 101-150 uw wide.
*043415: Azygospores are 151-200 wu wide.
*043416: Azygospores are 201-250 yu wide.
*043417: Azygospores are > 250 u wide.
Suspensors
008632: Two supporting cells (suspensors) are present.
*043418: Suspensors (supporting cells) are equal in size.
*043419: Both suspensors are spherical (globose).
*043420: Only the larger suspensor is spherical (globose).
Chlamydospores
008363: Chlamydospores are present.
008993: Chlamydospores occur singly.
008994: Chlamydospores are catenulate (in chains).
008996: Chlamydospores are spherical (length to breadth ratio
is 1.0-1.05).
*043421: Chlamydospores are prolate spheroidal (length to breadth
ratio is 1.06-1.15).
*043422: Chlamydospores are broadly ellipsoidal (length to breadth
ratio is 1.16-1.30). |
*043423: Chlamydospores are ellipsoidal (length to breadth ratio
is 1.31-1.6).
*043424: Chlamydospores are cylindrical.
008426: Chlamydospores are pigmented.
*043425: Chlamydospores are refringent (refractive).
*043426: Chlamydospore walls are smooth.
*043427: Chlamydospores are 0.1-1.0 “ wide.
*043428:
*043429:
*043430:
*043431:
*043432:
*043433:
*043434:
*043435:
*043436:
*043437:
*043438:
*043439:
*043440:
*043441:
*043442:
*043443:
*043444:
*043445:
*043446:
*043447:
*043448:
*043449:
*043450:
*043451:
*043452:
*043453:
*043454:
*043455:
*043456:
*043457:
Chlamydospores are 1.1-2.0 4 wide.
Chlamydospores are 2.1-3.0 u wide.
Chlamydospores are 3.1-4.0 u wide.
Chlamydospores are 4.1-5.0 u wide.
Chlamydospores are 5.1-6.0 u wide.
Chlamydospores are 6.1-7.0 « wide.
Chlamydospores are 7.1-8.0 u wide.
Chlamydospores are 8.1-9.0 uw wide.
Chlamydospores are 9.1-10 “ wide.
Chlamydospores are 11-15 wu wide.
Chlamydospores are 16-20 u wide.
Chlamydospores are 21-25 yu wide.
Chlamydospores are > 25 u wide.
Chlamydospores are 0.1-1.0 uw long.
Chlamydospores are 1.1-2.0 uw long.
Chlamydospores are 2.1-3.0 u long.
Chlamydospores are 3.1-4.0 u long.
Chlamydospores are 4.1-5.0 u long.
Chlamydospores are 5.1-6.0 u long.
Chlamydospores are 6.1-7.0 uw long.
Chlamydospores are 7.1-8.0 u long.
Chlamydospores are 8.1-9.0 uw long.
Chlamydospores are 9.1-10 u long.
Chlamydospores are 11-15 yu long.
Chlamydospores are 16-20 wu long.
Chlamydospores are 21-25 yu long.
Chlamydospores are 26-30 u long.
Chlamydospores are 31-35 yu long.
Chlamydospores are 36-40 yu long.
Chlamydospores are > 40 uw long.
Colony Pigmentation
173
020037: Nondiffusible pigment is produced.
020001: Colonies are pure (paper) white on solid medium.
020039: Nondiffusible brown pigment is produced.
020043: Nondiffusible golden (yellow) pigment is produced.
020002: Colonies are gray on solid medium.
174
Growth Temperatures
017032:
017012:
017013:
017066:
017037:
017014:
017033:
017052:
017034:
017074:
017043:
017017:
017018:
017019:
017020:
Growth occurs at 5°C.
Growth occurs at 10°C.
Growth occurs at 15°C.
Growth occurs at 17°C.
Growth occurs at 20°C.
Growth occurs at 25°C.
Growth occurs at 30°C.
Growth occurs at 33°C.
Growth occurs at 35°C.
Growth occurs at 36°C.
Growth occurs at 40°C.
Growth occurs at 45°C.
Growth occurs at 50°C.
Growth occurs at 55°C.
Growth occurs at 60°C.
Source of Isolation
002012: What was the specific source of isolation (e.g., kind of
water, soil, etc., species and organ and tissue of plant,
animal, etc.)?
Acknowledgements
This work was supported in part by National Science Foundation
Grant DIR89-15137 to SCJ.
The authors kindly thank Dr. Guozhong Ma for reviewing this paper.
175
REFERENCES
Alexopoulos, C.J. and Mims, C.W. 1979. Introductory Mycology.
John Wiley & Sons, New York, 632 pp.
Dabinett, P.E. and Wellman, A.M. 1973. Numerical taxonomy of the
genus Rhizopus. Can. J. Bot. 51:2053-2064.
Ehrenberg, C.G. 1820. De Mycetogenesi. Bonnae.
Ellis, J.J. 1985. Species and varieties in the Rhizopus arrhizus-
Rhizopus oryzae group as indicated by their DNA
complementarity. Mycologia 77:243-247.
Ellis, J.J. 1986. Species and varieties in the Rhizopus microsporus
group as indicated by their DNA complementarity. Mycologia
78:508-5 10.
Fischer, A. 1892. Phycomycetes. In Rabenhorst’s Kryptogamen-
flora, Die Pilze, Vol 4. Kummer, Leipzig, 505 pp.
Hanzawa, J. 1915. Studien uber einige Rhizopus-Arten. Mykol.
Zentralbl. 5:230-246.
Hawksworth, D.L., Sutton, B.C., and Ainsworth, G.C. 1983.
Ainsworth & Bisby’s dictionary of the fungi (including the
lichens), 7th ed. Commonwealth Mycological Institute, Kew,
Surrey, England, 445 pp.
Hesseltine, C.W. and Ellis, J.J. 1961. Notes on Mucorales, especially
Absidia. Mycologia 53:406-426.
Inui, T., Takeda, Y., and lizuka, H. 1965. Taxonomical studies on
genus Rhizopus. J. Gen. Appl. Microbiol. 11(suppl.):1-121.
Jong, S.-C., Davis, E.E., McManus, C., and Krichevsky, M.I. 1991.
Computer coding of strain features of the saprolegnian fungi.
Mycotaxon 41:407-418.
Jong, S.-C., Ho, H.H., McManus, C., and Krichevsky, M.I. 1989.
Computer coding of strain features of the genus Phytophthora.
Binary 1:187-193.
Jong, S.-C., Ho, H.H., McManus, C., and Krichevsky, M.I. 1992.
Computer coding of strain features of the genus Pythium.
Mycotaxon 44:301-314.
Jong, S.-C., Holloway, L., McManus, C., Krichevsky, M.I., and
Rogosa, M. 1988. Coding of strain features for computer-aided
identification of yeasts. Mycotaxon 31:207-219.
176
Lendner, A. 1908. Les Mucorinées de la Suisse. Matériaux pour la
flora cryptogamique Suisse. K.-J Wyss, Berne, 180 pp.
Liou, G.Y., Chen, C.C., Hsu, W.H., and Chien, C.Y. 1990. Atlas of
the genus Rhizopus and its allies. Mycological monograph No. 3.
Food Industry Research and Development Institute, Taiwan,
32 pp.
Naumov, N.A. 1939. Clés des Mucorinées. Encyclopédia
mycologique, Vol. 9. Lechevalier, Paris, pp. 1-137.
Rogosa, M., Krichevsky, M.I., and Colwell, R.R. 1971. Method for
coding data on microbial strains for computers (edition AB). Int.
J. Syst. Bacteriol. 21:1A-184A.
Rogosa, M., Krichevsky, M.I., and Colwell, R.R. 1986. Coding
microbiological data for computers. Springer-Verlag, New York,
299 pp.
Schipper, M.A.A. 1984. A revision of the genus Rhizopus. I. The
Rhizopus stolonifer-group and Rhizopus oryzae. Stud. Mycol.
25:1-19.
Schipper, M.A.A. and Stalpers, J.A. 1984. A revision of the genus
Rhizopus. WU. The Rhizopus microsporus-group. Stud. Mycol.
25:20-34.
Yang, G.F. and Jong, S.C. 1984 A new obligate azygosporic species.
Mycotaxon 20:397-340.
Zimmermann, O.E.R. 1871. Das genus Mucor. In Programm der
Realschule I. Ordnung zu Chemnitz.
Zycha, H. 1935. Pilze II. Mucorineae. In Kryptogamenflora der
Mark Brandenburg, Vol. 6a. Gebruder Borntraeger, Leipzig,
264 pp.
Zycha, H., Siepmann, R., and Linnemann, G. 1969. Mucorales.
Eine Beschreibung aller Gattungen und Arten _ dieser
Pilzgruppe. J. Cramer, Lehre, 355 pp.
MY COTAXON
Volume XLVI, pp. 177-182 April-June 1993
THREE NEW HYALOSCYPHACEOUS FUNGI
FROM NORWAY AND GREENLAND
SIGURD OLSEN *
Skarveien 7, N-8550 Lédingen, Norway
JOHN HAINES
Biological Survey, rm 3132 CEC, NYS Museum
Albany, NY 12230, U.S.A.
SIGMUND SIVERTSEN
UNIT, Botanical Department, N-7004 Trondeim, Norway
SUMMARY: Three species of Hyaloscyphaceae from Norway and Greenland,
Urceolella aasii sp. nov. on Urtica, Hyalopeziza archangelica sp. nov. on
Angelica/Heracleum, and Hyalopeziza groenlandica sp. nov. on Angelica, are
described and briefly discussed. Collections were made by the first author
unless otherwise stated.
KEY WORDS: Greenland; Hyaloscyphaceae; Norway
Urceolella aasii Olsen & Haines, sp. nov. Pig)
Apothecia diametro usque ad .5 mm, aurantio-ochracea. Pili usque ad 250 um longi, 6-
16 pm lati, non-septati, praecrassotunicati, 11 todo non colorati, tabida in 3% KOH.
Sporae 6.5-9.5 um longae. 1.0-1.8 um crassi. Ad caulis emortuis Urticae.
Holotypus: Norway, Nordland, Rana: Ytteren. On Urtica dioica. 22 V 1991, S. Olsen.
In herb BG.
Etymology: Named in honor of Norwegian mycologist Olav Aas.
Apothecia up to .5 mm diam., orange to ochraceous, seated on very short stipe ca.
2/3 the diam. of the cup, disc orange-ochraceous, exposed and concave when moist,
covered by hairs when dry. Excipulum thin-walled, buff-orange fextura angularis
composed of cells ca. 12 4m in diam., outer layer flattened, almost forming a cortex.
Subhymenium and inner cells of excipulum containing orange oil-globules in revived
exiccata. Hairs up to 250 wm by 6-16 wm diam., cylindric or tapered to a blunt point
at the apex, straight or occasionally slightly curved or forked, non-septate with a fine
*Sigurd Olsen died November 23, 1992
178
lumen 1-2 um wide except at the base where it is wider, hyaline, smooth, arising
perpendicularly from the margin or outer exciple, sometimes with a plate-like or
rooting base, flexible, not easily breaking, slowly and almost completely dissolving in
warm 3% KOH to leave only a ghost-like outline, unchanged in Melzer’s solution.
Some hairs agglutinated apically by hyaline amorphous matter which stains in cotton
blue. Hair walls and lumen unstained with cotton blue. Short cylindrical hairs with
hemispherical tips and apically enlarged lumen also present. Asci 35-45 (57) x 4.5-6.0
um, 8-spored, cylindrical or enlarged in the mid portion, subtended by a crozier at
least in early development, with a minute apical pore plug which turns bright blue in
Melzer’s solution. Spores 6.5-9.5 1.0-1.8 (2.5) um, non-septate, cylindric-fusiform,
slightly enlarged above, straight or very slightly curved, without conspicuous
inclusions. Paraphyses evenly cylindrical, 1.3-1.7 um wide level with the ascus tips in
the hymenium, containing yellowish oil globules, straight, unbranched except at the
base.
Host: Known only from moist previous years stems of Urtica dioica in Spring or
summer. The type material also contains Crocicreas, Leptosphaeria, and a pycnidial
fungus.
Distributions: Known only by three collections from near or above the Arctic Circle
in Norway.
Collections examined: NORWAY; Trons., Harstad, Sdrvik. 27 VI 1989 on Urtica dioica in
herb. BG; Nordland, Rana, Ytteren. 25 V 1990 on Urtica dioica in herbs. BG, NYS; See also type.
This small, herbicolous discomycete fits into the Hyaloscyphaceous genus
Urceolella as defined by Korf & Kohn (1980), with its long, thick-walled hairs that
dissolve in KOH and remain unstained in Melzer’s solution. It is distinct within
Urceolella with its orange to ochraceous pigmentation and long hairs up to 250 um
arising perpendicularly from the substrate. It appears to be closely related to U.
crispula, from which it is distinguished by its larger size, larger spores, and colored
disc as compared to the watery white disc of U. crispula. Raschle (1977) describes
U. crispula as white or light beige. Another, perhaps even more closely related,
though isolated species, U. tuberculiformis, has the same type of hairs with T-like
bases. This species, as described by Dennis (1963), differs in having no pigmentation,
although Raschle (1977) describes it as yellowish white with a thick-walled yellow
excipulum, and asci, ca. 70 x 10 wm, longer and broader spores (10-12 x 2.5-3.0 um),
and growing on Aquilegia in the USA and Adenostyles in Italy.
Figs. 1-5. 1. Urceolella aasii, Hairs, asci, paraphyses, and spores from Paratype,
S6rvik. 2-4. Hyalopeziza archangelica. 2. Asci, paraphyses, hairs, and spores from
holotype. 3. Spores and hairs from paratype, Trondheim. 4. Spores from paratype
specimen, Skanland. 5. Hyalopeziza groenlandica Ascus, paraphyses, hairs, and spores
from holotype.
180
Hyalopeziza archangelica Olsen & Sivertsen, sp. nov. Fig. 2-4
Apothecia cupulata, sessilia, ad 1 mm diametro, rose, arcte setulosa. Pili hyalini, 25-65
uum longis, basi ad 6 um crassi, solidis. Asci 70-110 um longis, 15-17 4m crassi, in Iodo
non colorati. Paraphysis cylinribus. Sporae 10-15 (18.5) um longis, 4.0-6.5 (-7.0) pm
crassi, 0-3-septatae. Ad caulis emortuis Angelica et Heracleum.
Holotypus: Norway, Nordland, Rana, Fisktj6nna. On Angelica archangelica, 26 V
1990. Olsen & Sivertsen. In herb. BG.
Etymology: Named for its substrate, Angelica archangelica.
Apothecia sessile, cupulate, up to 1 mm across, mostly smaller, gregarious, whitish to
pink. Rim and flanks set with scattered, short hairs. Excipulum of thick-walled,
hyaline, angular, + isodiametric cells 6, 5-10 wm across. Hairs 25-70 ym. long,
refractive, solid above with a lumen at the base, + straight, rather evenly tapering
towards an acute apex. Not loosing glassiness in warm 10% KOH and not
colored by Meltzer’s or Congo Red. Asci 70-110, 15-17 um thin-walled, some apical
thickening when immature, 8-spored, arising from a crozier. Spores 10-15 (18.5) x
4-6 (7) um, hyaline, smooth, 0-2-, mostly 1-septate, often with prominent oil-globules.
Paraphyses cylindrical, 2.0-2.8 jm thick, with rounded apex, sparingly septate,
sometimes branched above the middle, not overtopping the asci, scarce.
Host: On basal parts of previous years stems of Angelica and Heracleum.
Distribution: Probably widespread over the northern hemisphere, but overlooked
because of its small size and nondistinctive color. A specimen cited by Korf (1980)
and presumed to be the same, is from Finmark, Norway, but it has not been
examined by the authors.
Collections examined: NORWAY, Nordland; Rana, Fisktjonna, 20 V 1991 on Angelica, S.
Olsen, herbarium BG; ‘Tromso, Skdnland, near airport, 11 VI 1991 on Heracleum, S. Olsen,
herbarium BG; Sér-Trondelag, Trondheim, Lillegardsbakken, III 1992, on Heracleum, S. Sivertsen,
herbarium BG; GREENLAND, Kuanit, east of Godhavn, on Angelica, P. Milan Petersen, herbarium
CUP as 5946; Also see type.
This is the same fungus from Finmark and Greenland as tentatively refered
to as "Chaetonaevia archangelica" by Korf (1980). Korf clearly did not intend to
legitimize the name according to the code in that publication and since it has so many
similarities to Hyalopeziza, it will be described here as a Hyalopeziza in subgenus
Unguicularia as defined by Korf & Kohn (1980). Its hairs are unchanged by KOH,
and the lumen is present only at the base. The excipulum is composed of +
isodiametric angular cells, which, according to Huhtinen (1987) is acceptable in the
genus.
The species seems to be closely related to Hyalopeziza raripila (H6hn.)
Huhtinen, but differs in its larger size, different color, longer hairs, asci, and spores
which are septate at maturity.
181
Hyalopeziza groenlandica Olsen, sp. nov. Fig. 5
Apothecia gregaria, cupulata, ad 0.5 mm lata, sessilia, ochraceo-rosei. Pili hyalini, ad
30 um longis, in apicem solidis attenuatis. Asci 55-60 4m longis, octosponi, poro in Todi
non colorati. Paraphyses cylindracea, apicem solidis attenuatis. Sporae 15-20 x 2.8-4.0
uum. Ad caulis emortuis Angelicae.
Holotypus: Greenland, Kuanit, approx. 4 km. east of Godhavn. Leg. P. Milan
Petersen 5 VIII 1970 in herbarium CUP (as CUP 59588).
Etymology: For the type locality, Greenland.
Apothecia yellowish-pink, sessile, cupulate, gregarious, up to 0.5 mm across. Margin
with short, hyaline hairs. Excipulum of slightly thick-walled, elongate, hyaline cells.
Hairs 25-35 um long, refractive, tapering, irregularly flexuose, hooked apically,
straight or sharply bent basally, with lumen at the base, refractivity not lost in warm
10% KOH, without color change in Meltzer’s solution. Asci 55-60 um long,
somewhat thick-walled, 8-spored, without croziers, pore not blued in Melzer’s solution
even with KOH pretreatment. Spores 15-20 x 2.8-4.0 um, wide fusiform, slightly
curved, aseptate, aguttulate. Paraphyses numerous, with irregular refractive, apical
protuberances, unchanged in KOH or Melzer’s, slightly exceeding the asci.
Host: Previous year’s stems of Angelica.
Distribution: So far known only from the type collection.
Korf (1980) tentatively assigned the name "Laetinaevia archangelica" to this
collection, but he was clear in not formally describing it as a new species. The first
author is inclined to assign this species to Hyalopeziza subgenus Unguiculella (sensu
strictu) Korf & Kohn (1980) on the basis of its glassy hairs and paraphyses. This
combination requires that a species name other than Korf’s be chosen as the epithet
"archangelica" is occupied by the new species Hyalopeziza archangelica Olsen &
Sivertsen described above. Korf stresses the paraphyses, which he described as
"coiled" and, although the general appearance is very much the same as H.
_archangelica Olsen & Sivertsen it differs in the glassy protuberances of the
paraphyses, as well as its quite different spores.
182
ACKNOWLEDGEMENTS
The authors wish to thank Dr. Linda Kohn for her prepublication review of
this work.
LITERATURE CITED
Dennis, R. W. G. 1963. A redisposition of some Fungi ascribed to the
Hyaloscyphaceae. Kew Bulletin 17: 319-379.
Huhtinen, S. 1987 (1988). Five glassy-haired Hyaloscyphaceae. Karstenia 27: 8-14.
Korf, R. P. 1980. Inoperculate discomycetes of the arctic and alpine zones of
Finmark, Lapland, and Greenland. In G. Laursen & J. Ammirati. Arctic and
Alpine Mycology I: 27-37.
Korf, R. P. & Linda M. Kohn 1980. Revisionary studies in the Hyaloscyphaceae. I.
On Genera with "glassy" hairs. Mycotaxon 10: 503-512.
Raschle, P. 1977. Taxonomische Untersuchungen and Ascomyceten aus der Familie
der Hyaloscyphaceae Nannfeldt. Sydowia 29: 170-236.
MY COTAXON
Volume XLVI, pp. 183-192 April-June 1993
TAXONOMIC HISTORY
OF THE OOMYCETE GENUS
THRAUSTOTHECA
WILL H. BLACKWELL AND MARTHA J. POWELL
Department of Botany, Miami University, Oxford, Ohio
45056
ABSTRACT
The nomenclature of Thraustotheca(Saprolegniales)
is reviewed. The rationale for the proper citation of
authorship of TT. clavata as (de Bary ex Biisgen)
Humphrey is explained. It is concluded that at present
only two validly published or adequately described
species of Thraustotheca_ should be recognized, T.
clavata and T. primoachlya. The inclusion of the
latter taxon in the genus is questioned based on current
consideration of primary sporangial characters as most
significant in generic delimitations.
Many genera of Oomycetes need contemporary
taxonomic and nomenclatural review. Sparrow’s 1960 review
of Oomycete genera in his monumental work on
Phycomycetes is the most recent extensive synoptic treatment
of the family Saprolegniaceae. Dick (1973) provided a list of
genera and species of the Saprolegniaceae in chronological
order of their descriptions. New species of the family have
been described in more recent years, yet within the last forty
years the only genera of Saprolegniaceae which have been
monographed are Achlya (Johnson, 1956), Aphanomyces
(Scott, 1961) and Saprolegnia (Seymour, 1970). In addition
184
Langsam (1986, 1987) has carried-out revisionary
investigations on the genus Brevilegnia . In the following
account we review the current taxonomic state of one genus of
the Saprolegniaceae, Thraustotheca Humphrey. We clarify
nomenclatural errors and identify taxa which require
revisionary study.
History and Authorship of Thraustotheca clavata
The original species of the genus Thraustotheca
Humphrey (1893), 7. clavata , is based on the taxon
Dictyuchus clavatus. Authorship of the epithet c/avatus is
usually credited (Coker and Hyman, 1912; Coker, 1923; Coker
and Matthews, 1937; Shanor, 1937; Wolf, 1944; Sparrow,
1960) solely to Anton de Bary (1888), although some also
noted (Coker and Hyman, 1912; Weston, 1918; Coker and
Matthews, 1937; Shanor, 1937) that Busgen (1882) first
incidentally published the species (i.e., D. clavatus ).
Busgen’s (1882) publication on sporangial and spore
formation provided an extensive description of asexual
development of D. clavatus , and as well an account of sexual
structures of the organism, in effect a complete description.
However, Busgen credited the species to de Bary, who had
received it in 1880 as part of a mixed algal collection obtained
by E. Stahl at Vendenheim, Germany. A preferred citation of
authorship of Dictyuchus clavatus would thus be de Bary ex
Biisgen (1882) or simply Busgen (cf. Article 46.3,
CURSE Code of Botanical Nomenclature, Grueter et al.,
1988).
De Bary mentioned Dictyuchus clavatus in his 1884
text without description, referencing his student’s (i.e., M.
Busgen’s) dissertation. De Bary’s 1888 actual description of
the taxon provided additional morphological information, but
this is not relevant to the point of its valid publication. In fact,
de Bary’s 1888 publication was a posthumous paper edited by
Solms-Laubach (Weston, 1918), and it is difficult to ascertain
185
de Bary’s intent as to the description of D. clavatus as a new
species. Regardless, a proper citation of authorship under
Thraustotheca would be T. clavatus (de Bary ex Busgen)
Humphrey (1893), as listed in Farr et al. (1979).
Concept of the Genus Thraustotheca
Solms-Laubach (in de Bary, 1888) first noted the
possible generic distinction of D. clavatus within the
Saprolegniales; this was reiterated by Fischer (1892).
Humphrey subsequently (1893) described the genus
Thraustotheca based on this taxon, but without benefit of
living material. Whereas Saccardo (1899) cited the date of
publication of the generic name as 1892, Clements and Shear
(1931) cited it as 1893. This discordance over date of
publication is resolved by understanding that Humphrey
read his manuscript on Saprolegniaceae of the United States
before the American Philosophical Society in November of
1892, but the Proceedings were published in 1893, the actual
year of effective publication of the name. Coker and Hyman
(1912) clearly stated the morphological delimitation of
Thraustotheca (i.e., T. clavata). Over the years the generic
concept of Thraustotheca , however, has evolved (Coker and
Couch, 1923, 1924; Coker and Braxton, 1926; Coker, 1927;
Coker and Matthews, 1937; Wolf, 1944; Sparrow, 1960) , and
we can summarize it now as:
those Saprolegniaceae producing a thraustothecoid
sporangium (=generally club-shaped or subcylindric
with an inoperculate, quickly disintegrating or
rupturing sporangial wall), which contains no
dictyuchoid network but which usually contains
numerous primary cystospores (i.e., primary motile
stage suppressed); oogonia with multiple oospores;
and hyphae with relatively broad diameters which
form a spreading hyphal mass.
186
These characters, in combination and in predominance,
distinguish Thraustotheca from other genera of the
Saprolegniaceae which are often considered morphologically
similar: e.g. Achlya, Brevilegnia, Calyptralegnia, Dictyuchus,
and Geolegnia.
Setting generic limits within the Saprolegniaceae is
sometimes problematic because of morphological plasticity.
Some genera exhibit more than one type of sporangial
discharge, yet it is generally accepted that the mechanism of
sporangial discharge is a primary character distinguishing
genera. For example, one species of Thraustotheca recognized
(T. primoachlya ) does not exclusively produce sporangia with
thraustothecoid type of spore discharge. Decisions on generic
placements have traditionally been based on which sporangial
type predominates (Coker, 1923). Instead, more recently, the
type of spore discharge found in first-formed sporangia
(Langsam, 1986) has been used for generic assignment.
Accordingly, Langsam (1986) transferred Brevilegnia bispora
to Achlya because of achlyoid spore discharge of primary
sporangia and minisporangia. These viewpoints (i.e, relative
importance of predominant versus initial sprorangial type)
caution that clearer delimitation of taxa requires
developmental studies of isolates grown under varying
environmental conditions, as well as numerical comparison of
genera using all putatively distinguishing generic
characteristics.
Nomenclatural Summary
Several taxa (in addition to 7. clavata) have been
described within Thraustotheca , two of these (T. achlyoides
and TJ. unisperma ) eventually constituting bases for
recognition of new genera (respectively Calyptralegnia and
Brevilegnia) of Oomycetes. Following is a chronological
nomenclatural summary of names published as taxa of
Thraustotheca. Names in boldface are presently construed as
valid taxa of the genus.
187
Thraustotheca Humphrey, 1893
Thraustotheca clavata (de Bary ex Biisgen)
Humphrey, 1893.
The basionym is Dictyuchus clavatus de Bary ex
Biisgen, 1882. Citation as D. clavatus Busgen is
equally permissible (Article 46.3, ICBN).
Thraustotheca caucasica Voronov, 1922. This name has
been overlooked by virtually all authors. It was,
however, noted in 1969 in Index of Fungi, a Supplement
to Petrak’s Lists. Although validly published, T.
caucasica must be considered a doubtful or exluded
taxon. Though seemingly similar to T. primoachlya ,
e.g., possessing tuberculate oogonia, there is insufficient
descriptive information (including the lack of an
illustration, as well as any information on antheridia) to
make a positive determination of identity. Should it be
possible to determine that it is the same as T.
primoachlya , the epithet caucasica would assume
priority.
Thraustotheca achlyoides Coker and Couch, 1923. =
Calyptralegnia achlyoides (Coker and Couch) Coker,
1927.
Thraustotheca primoachlya Coker and Couch,
1924.
Although we are recognizing J. primoachlya herein,
further assessment of this taxon as a species of
Thraustotheca is needed. In contrast to T. clavata, this
species has androgynous antheridial branches and
primary sporangia of the achlyoid type (producing the
usually predominating thraustothecoid type sporangia
later).
188
Thraustotheca unisperma Coker and Braxton, 1926. =
Brevilegnia unisperma (Coker and Braxton) Coker and
Couch in Coker, 1927. This is tantamount to transfer of
Thraustotheca unisperma var. unisperma, an autonym
created when var. /itoralis (see below) was described
(Coker and Braxton, 1926).
Thraustotheca unisperma var litoralis Coker and
Braxton, 1926. = Brevilegnia unisperma var. litoralis
(Coker and Braxton) Coker and Couch in Coker, 1927.
In Coker’s work, the transfer to Brevilegnia is
erroneously credited to Coker and Braxton. Because of
similarities of sporangial structure and dehiscence,
Salvin (1942) seemingly proposed that Brevilegnia be
only a subgenus of Thraustotheca , and hence that all
taxa described in or transferred to Brevilegnia should
be returned toZhraustotheca. Based on total
morphology (including one-spored oogonia), we favor
retention of Brevilegnia as a separate genus, and do not
favor such transfer of taxa back to Thraustotheca. We
note that Langsam (1987), though excluding certain
species, nonetheless retained the genus Brevilegnia .
Thraustotheca irregularis Coker and Ward in Ward,
1939. This isa nomen invalidum, lacking a Latin
diagnosis. Should it become desirable to recognize this
taxon, it would be necessary to provide such descriptive
information in Latin. Dick (1973) questioned the status
of T. irregularis. Johnson (1956) considered the taxon
possibly similar to Achlya dubia Coker (1923), in effect
questioning its acceptance as a species.
Two taxa, thus, are presently recognized as valid species
of Thraustotheca, T. clavata and T. primoachlya.
This is in fact the composition of the genus arrived at de
facto by Coker (1927) when T. achlyoides and T. unisperma
were transferred out of the genus. This circumscription
189
recognizes that T. primoachlya does not exclusively produce
thraustothecoid sporangia, and hence at this time we are
following Coker’s (1923) conservative concept of
predominance of sporangial discharge for generic assignment.
Thraustotheca clavata shows only thraustothecoid sporangia,
whereas 7. primoachlya demonstrates a progression from
achlyoid to thraustothecoid sporangial structures. Since
Achlya dubia Coker , like T. primoachlya , can produce both
achlyoid and thraustothecoid types of sporangia, both
taxa should be _ investigated further. In fact, based on
diclinous antheridia and primary sporangia of the
thraustothecoid type, Achlya dubia may be more similar to T.
clavata than is T. primoachlya.. Revisionary studies on
species of Achlya and Thraustotheca with multiple types of
sporangial discharge will have to be conducted to decide
the validity of including species which demonstrate achlyoid
sporangial discharge, even if only for a limited duration of the
asexual cycle, within Thraustotheca. Central to the issue is
whether primary sporangial type should take precedence over
predominance of sporangial type. The dilemma is complex
because temperature may influence the relative ratio of each
type produced (Alabi, 1972). The current study is
foundational for future investigations.
Key to Recognized Species of Thraustotheca
1. Sporangia all thraustothecoid, club-shaped; oogonia
smooth; antheridia diclinous.......... T. clavata
2. Early sporangia achlyoid, later ones thraustothecoid,
often more elongate; oogonia with blunt projections;
antheridia androgynou................ T. primoachlya
190
ACKNOWLEDGEMENTS
We are indebted to the following persons for their
helpful and critical review of this manuscript: Dr. Deborah
M. Langsam, Department of Biology, The University of North
Carolina at Charlotte; Dr. Michael A. Vincent, Department of
Botany, Miami University; and Dr. David E. Padgett,
Department of Biological Sciences, The University of North
Carolina at Wilmington. This work was supported by a grant
(BSR 91-07451) from the National Science Foundation.
LITERATURE CITED
ALABI, R.O. 1972. Thraustothecoid sporangium formation
in three Saprolegniaceae from Nigeria. Trans. Brit.
_. Mycol. Soc. 58: 519-521.
BUSGEN, M._ 1882. Die Entwicklung der
Phycomycetensporangien. Jahrb. Wiss. Bot. 13: 253-285.
CLEMENTS, F.E. and C.L. SHEAR. 1931. The Genera of
Fungi. The H.W. Wilson Co., New York. iv + 496 p. and
58 pits.
COKER, W.C. 1923. The Saprolegniaceae, with Notes on
Other Water Molds. Univ. of North Carolina Press,
Chapel Hill. 201p. and 63 pits.
COKER, W.C. 1927. Other water molds from soil. J. Elisha
Mitchell Sci. Soc. 42: 207-226.
COKER, W.C. and H.H. BRAXTON. 1926. New water
molds from soil. J. Elisha Mitchell Sci. Soc. 42: 139-149.
COKER, W.C. and J.N. COUCH. 1923. A new species of
Thraustotheca. J. Elisha Mitchell Sci. Soc. 39: 112-114.
COKER, W.C. and J.N. COUCH. 1924. Revision of the
genus Jhraustotheca , with a description of a new species.
J. Elisha Mitchell Sci. Soc. 40: 197-201.
COKER, W.C. and O.W. HYMAN. 1912. Thraustotheca
clavata. Mycologia 4: 87-90.
COKER, W.C. and V.D. MATTHEWS. 1937.
Saprolegniales. North America Flora 2: 15-67.
191
DE BARY, A. 1884. Vergleichende Morphologie und
Biologie der Pilze, Mycetozoen und Bacterien. Wilhelm
Engelmann, Leipzig. xvi + 558 p.
DE BARY, A. 1888. Species der Saprolegnieen. Bot. Zeit.
46: 597-610, 613-621, 629-636, 645-653. (Paper edited
posthumously by H.G. Solms-Laubach. )
DICK, M.W. 1973. Saprolegniales. IN: The Fungi, An
Advanced Treatise. Ainsworth, G.C., F.K. Sparrow, and
A.S. Sussman, eds. Academic Press, New York and
London. pgs. 113-144.
FARR, E.R., J.A. LEUSSINK and F.A. STAFLEU (eds.).
1979. Index Nominum Genericorum (Plantarum), Vol.
3. Reg. Veg. 102: 1277-1896.
FISCHER, A. 1892. Saprolegninae. IN: Rabenhorst’s
Kryptogamenflora 1(4): 310-383.
GREUTER, W., H.M. BURDET, W.G. CHALONER, V.
DEMOULIN, R. GROLLE, D.L. HAWKSWORTH, D.H.
NICOLSON, P.C. SILVA, F.A. STAFLEU, E.G. VOSS
and J. McNEILL. 1988. International Code of Botanical
Nomenclature (ICBN) . Adopted by the Fourteenth
International Botanical Congress, Berlin, July-August,
1987. Regnum Vegetabile 118: xiv + 328 p.
HUMPHREY, J.E. 1893. The Saprolegniaceae of the United
States, with notes on other species. Trans. Amer. Phil.
Soc. (N.S.) 17: 63-148.
INDEX OF FUNGI. 1969. A Supplement to Petrak’s Lists.
1920-1939. Commonwealth Mycological Institute, Kew,
Surrey. p. 153. (citation source of T. caucasica ).
JOHNSON, T.W. 1956. The genus Achlya: Morphology and
Taxonomy. Univ. Michigan Press, Ann Arbor. xv +
180 p.
LANGSAM, D.M. 1986. Achlya bispora: A taxonomic
reassessment of Brevilegnia bispora . Mycologia 78: 600-
604.
LANGSAM, D.M. 1987. Notes on the genus Brevilegnia:
Two exluded taxa. Mycologia 79: 323-324.
SACCARDO, P.A. 1899. Sylloge Fungorum 14: 36.
192
SALVIN, S.B. 1942. Variations of specific and varietal
character induced in an isolate of Brevilegnia.
Mycologia 34: 38-51.
SCOTT, W.W. 1961. A monograph of the genus
Aphanomyces. Virginia Agri. Exp. Sta. Tech. Bull. 151:
1-95.
SEYMOUR, R.L. 1970. The genus Saprolegnia. Nova
Hedwigia 19: iv -124.
SHANOR, L. 1937. Observations on the development and
cytology of the sexual organs of Thraustotheca clavata (de
Bary) Humph. J. Elisha Mitchell Sci. Soc. 53: 119-136.
SPARROW, F.K. 1960. Aquatic Phycomycetes, 2nd. rev. ed.
Univ. Michigan Press, Ann Arbor. xxv + 1187 p.
VORONOV, I. 1922. Thraustotheca caucasica. Mater.
Mikol. Fitopat. Ros. 4(1): 57-58.
WARD, M.W. 1939. Observations on a new species of
Thraustotheca. J. Elisha Mitchell Sci. Soc. 55: 346-
352.
WESTON, W.H. 1918. The development of Thraustotheca, a
peculiar water-mould. Ann. Bot. 32: 155-173.
WOLF, F.T. 1944. The Aquatic Oomycetes of Wisconsin.
Part 1. Univ. Wisconsin Press, Madison. 64 p.
MYCOTAXON
Volume XLVI, pp. 193-199 April-June 1993
BIOSTATISTICAL CHARACTERIZATION OF THE GENUS OIDIUM
MARCO T. IALONGO
Istituto Sperimentale per la Patologia Vegetale,
Via C.G. Bertero 22, Rome, Italy
A method that takes advantage of biostatistical analysis is proposed for differentiating
between species of the genus Oidium (anamorph of Erysiphoideae), since it is often difficult to
distinguish one from the other using the traditional parameters. In addition to the usual
parameters, new quantitative morphological parameters are introduced: area, circumference,
maximum diameter, shape factor of the circumference, shape factor of the area of the
appresoria; true length of conidial foot-cell; area, circumference, shape factor of circumference,
shape factor of the area of the conidia. Measurements were made of the anamorphs of various
species of Erysiphaceae selected from those that have indistinguishable forms according to the
traditional measurement methods. The series of values obtained for the single parameters were
compared two by two (the values of each sample with the corresponding values of all the
others), calculating Student’s t for each comparison. The average linkage cluster analysis was
then applied, and the relative dendrogram constructed. The latter demonstrates how the
anamorphs most similar to one another are found on the plants systematically most like one
another, and so on. Apart from demonstrating the possibility to differentiate between anamorphs
that would otherwise be indistinguishable, the strict adherence of the conidial form of these
obligate pathogens to the relative matrix is also shown. This method is useful both during
epidemiological surveys, thus permitting suitable chemical intervention, as well as during
systematic investigations on the pathogens under study. The standardization of the method,
proposed for the first time in the field of powdery mildews, begins with the considered
anamorphs being distributed in a sequence of six-figure codes (given the name "morphotype")
expressing the following six groups of characteristics: type of appressoria, conidial shape,
placement of conidia, absence or presence of fibrosin bodies, conidial outline, shape of
conidiophore foot-cell. The biostatistical analysis described above will successively be applied to
the anamorphs of each single morphotype.
Introduction and Aims
As is known, in the subfamily Erysiphoideae of the family Erysiphaceae, the
teleomorphs of the various species can be fairly straightforwardly differentiated.
The same cannot be said for the anamorphs (belonging to the genus Oidium
Link), which, in the light of what is currently known, are often morphologically
indistinguishable from each other. Furthermore, the habit of using the name of
the teleomorph to indicate the anamorphs as well has more than ever made
people refrain from carrying out a close study of the latter. Commenting on the
lack of literature on the subject, Braun has recently published a few
contributions (1980, 1982, 1982a).
194
In addition, since several powdery mildews are found, in nature, at the
anamorphic state and infrequently, or at least, not always, develop into the
perfect form, it has been considered worthwhile to make an accurate
investigation of this conidial form.
However, a deeper study of the individual anamorphs is not just an
attempt to answer taxonomic queries; the possibility of differentiating one form
from another also leads to a more thorough understanding of the epidemiology
of the individual species and thus knowing the likelihood, or not, of the
mildews on spontaneous plants infecting agricultural crops or spreading among
them. The advantages to the planning of control methods are obvious.
The aim of the present work is to reach such results, proposing a
methodology based on biostatistical analysis. The analysis is carried out using
both classical and new quantitative morphological parameters. The research on
new parameters is even more necessary for the anamorphs whose descriptions
(when based only on the parameters usually adopted) do not permit exact
attribution to one species rather than another.
Materials and Methods
Five samples of "powdery mildews" were selected in the conidial state,
according to the following criteria.
As the first criterion, five anamorphs were chosen that, on the basis of the
usual characteristics considered in the description of an Oidium, presented an
identical "profile", 1.e.,
- type of appressoria: multi-lobed (type C of Boesewinkel, 1977);
- conidial shape: cylindrical;
- conidial placement: carried singly by the conidiophore (Pseudoidium-type
anamorph: cfr. Blumer, 1967);
- fibrosin bodies: inconspicuous;
- conidial outline: curvilinear;
- shape of the foot-cells of the conidiophore: cylindrical, flexuous at the
base.
Thus, according to the above characteristics, no differentiation exists
between the five samples.
As already mentioned, such a criterion was used to select five conidial
forms of Erysiphaceae, which are cosmopolitan on the following host plants:
1&2) Erysiphe convolvuli DC. on two hosts of the same species: Calystegia
sylvatica (Kit.) Griseb. - Fam. Convolvulaceae, but collected in areas far
from one another (this pair is witness to the effectiveness of the
proposed method);
3) The same E. convolvuli collected on a plant belonging to the family of 1
& 2), but of another genus - Convolvulus arvensis L.;
195
4) Another powdery mildew of a different genus than the preceding one -
Microsphaera trifoli (Grev. ) U. Braun on a host, Trifolium pratense L.,
belonging again to a different family (Leguminosae);
5) Another powdery mildew of the same genus as above but belonging to
another species - Microsphaera euonymi-japonici Viennot-Bourgin, on a
host of a family that is very different from the two preceding ones -
Euonymus japonicus L. fil. (fam. Celastraceae).
The identification of the first four conidial forms was confirmed also
with the respective teleomorphs.
All five samples were collected on the same day and (obviously with the
exception of one of the two Calystegia) in the same area, from plants found on
the outskirts of Rome that had not undergone any chemical treatments.
The measurements of the individual parameters of the anamorphs under
study were Statistically processed using a Kontron MOP. Videoplan image
analyzer connected via a telecamera to a Zeiss photomicroscope.
The quantitative characteristics considered for each sample are relative to
the following parameters:
1) area (projection surface) of the appressoria;
2) circumference (projection outline) of the appressoria;
3) maximum diameter (of the projection surface) of the appressoria;
4) shape factor of the circumference of the appressoria, based on the ratio
area/circumference (used to define regular or irregular structures: for
ellipses and irregular structures its value remains less than 1, while it
reaches unity in the case of a circular structure);
Dy) shape factor of the appressorium area, based on the minimum and
maximum area/diameter ratio (as above: in the case of circular
structures its value is 1, for irregular structures it remains less than
unity);
6) real length of basal cell of the conidiophore;
7) area (projection surface) of conidia;
8) circumference (projection outline) of conidia;
9) shape factor of the conidial circumference (see above);
10) shape factor of the conidial area (see above);
11) conidial length;
12) maximum width of conidia;
13) ratio length/maximum width of conidia.
From a research of the literature, the type of processing reported above
has never been previously applied to Erysiphaceae, in general.
196
A broad outline for statistical processing has, at most, been inferred by
Zar (1974) and Nordio (1977).
The series of values obtained for the individual parameters (totalling 13
series for each sample) have been compared two by two (the series from each
sample with the corresponding series of all the others), calculating Student’s t
for each comparison. In order to visualize the ratio between the above series,
the average linkage cluster analysis was applied to the symmetrical matrix of t,
to allow the construction of the relative dendrogram.
Standardization of the Method
As already explained, in order to process the data collected, the
preliminary condition was imposed that the anamorphs to be compared belong
to the same "profile", based on six characteristics (obviously it is not necessary
to differentiate between two or more anamorphs if they already belong to
different profiles).
To standardize this sequence of descriptive characteristics, for which the
term "morphotype", as used by Ainsworth, 1971, is proposed, it is considered
convenient to have a coding.
The morphotype will be characterized by numbers chosen in the six
figures, always in the same succession; one then proceeds with the following
coding (Fig. according to Boesewinkel (loc. cit) and Braun (1987)).
I-FIGURE Type of appressoria
Indistinct: Boesewinkel, fig. 3E, Braun, Fig. 1a.
Nipple-shaped: Boesewinkel, fig. 3A, Braun, Fig. 1b-g.
Slightly or moderately lobed: Boesewinkel, Fig. 3D, Braun, fig. 1h.
Multi-lobed: Boesewinkel, fig. 3C, Braun, Fig. 11.
II-FIGURE Conidial shape
Sashes
As is known, conidia of the genus Oidium can assume very different
shapes, which basically lead back to three:
1. _Ellisoid or ovoid: Braun, Figs. 3A and 3B.
2. Cylindric (when the two largest sides are parallel for a long tract: Braun,
Fig. 3C).
3. Botuliform or doliform (with nonparallel sides and with the two ends
prominent: Braun, Fig. 3D).
III-FIGURE Placement of the conidia
—
Formed singly.
2. Produced in chains.
IV-FIGURE Absence or presence of fibrosin bodies
1. Present - well-developed fibrosin bodies.
2. Absent - inconspicuous fibrosin bodies.
197
V-FIGURE Conidial outline
1. Curvilinear.
2. Angular, as for the case of Sawadaea.
VI-FIGURE Shape of conidiophore foot-cell
This is also very variable but lead back to seven basic morphologies:
Gradually decreasing in diameter from base to top: Boesewinkel, Fig. SE.
Cylindrical (straight, cylindrical foot-cell: Boesewinkel, Fig. 5A).
Gradually decreasing in diameter from top to base: Boesewinkel, Fig. 5D.
Swollen at the base.
Moniliform (swollen foot-cell with a conspicuously enlarged basal area:
Boesewinkel, Fig. 5B; typical of anamorphs of Blumeria graminis (DC.)
Speer).
6. Flexuous.
7. Conidiophore curved in the basal half: Braun, Fig. 65c.
A cat ad dae
As an example of what is listed above, the morphotype under study is: 4 2
1 2 1 6, and so on.
Thus, in order to recognize and differentiate these anamorphs, one first
determines the respective morphotypes, grouping together those that are
identical. Then, the anamorphs of each morphotype are biostatistically analyzed
as described above, with the aim of further differentiating one from the other.
Results
The dendrogram constructed with the processed data is given in Fig. 1. It
shows how the measurements that are most similar to one another belong to
the two Enrysiphe convolvuli on the two samples of Calystegia. Then, with
increasing order of t, it can be seen that this pair is close to the same E.
convolvuli collected on the other convolvulacea, but well differentiated from it.
The measurements on the powdery mildews of the other two _ botanical
families are considerably dissimilar from the preceding measurements.
Finally, by calculating, for each of the three families under study, the
overall average of their Student’s t, we can note how a closer similarity exists
between the leguminosa and the three convolvulaceae than between the latter
and the celastracea: this is in agreement with the similarity between the
botanical families as reported by Hutchinson (1969).
Using the methodology applied here, we can obtain the differentiation of
the anamorphs under study, which would be impossible to do by observation
only.
- Moreover, the biostatistical analysis used on the individual samples leads
to a deeper insight into the morphological characterization of the various
anamorphs: this information constitutes a valuable databank that can also be
used for systematic analyses of the genus Oidium.
198
Euonymus japonicus
Convolvulus arvensis
Trifolium pratense
S
=
3s
>
aS}
Ry
&
S
—| Calystegia sylvatica
2.0
3.0
4.0
Se
6.0
7.0
8.0
Fig. 1 Cluster analysis dendrogram for single linkage
applied to the symmetrical matrix of Student’s t.
199
Discussion and Conclusions
The methodology proposed in this work has enabled to distinguish the
positions of reciprocal similarity between the five anamorphs examined. The
Statistical analysis has outlined both a strict adherence of the conidial shape of
the pathogen to its host, as well as its precise relationship to the relative
teleomorph.
The results confirm the well-known determining influence of the host
plants, a true and proper matrix, on at least the anamorphic morphology of
the fungus; thus, it can be utilized to investigate the natural distribution of such
obligate pathogens.
In the meanwhile the method provides valuable information to enable a
deeper insight into the systematics of Erisifaceae.
It can be concluded that, at least in the case of anamorphs, the method
presented appears to justify the authors who have proposed a classification of
these pathogens that strictly adheres to the taxonomic position in the botanic
system of the respective host plants.
The author wishes to thank Prof. A. Rambelli for kindly reviewing the
manuscript.
References
Ainsworth, G.C. (1971). Aisnworth & Bisby’s Dictionary of the Fungi.
Commonwealth Mycological Institute, Kew, Surrey, 663 pp.
Blumer, S. (1967). Echte Mehltaupilse (Erysiphaceae). Gustave Fischer Verlga,
Jena, 436, pp.
Boesewinkel, H.J. (1977). Identification of Enrysiphaceae by _ conidial
characteristics. Revue Mycol., 41: 493-507.
Braun, U. (1980). Morphological studies in the genus Oidium. Flora, 170:
77-90.
Braun, U. (1982). Morphological studies in the genus Oidium (II). Zbl.
Mikrobiol., 137: 138-152.
Braun, U. (1982a). Morphological studies in the genus Oidium (III). Zbl.
Mikrobiol., 137: 314-324.
Braun, U. (1987). A monograph of the Erysiphales (powdery mildews). Nova
Hedwigia, beiheft 89. J. Cramer, Berlin-Stuttgart, 1-700.
Hutchinson, J. (1969). Evolution and phylogeny of flowering plants. Academic
Press, London and New York, 717 pp.
Nordio, V. (1977). Festuca alpestris R. et S. nelle prealpi bellunesi. Giorn. Bot.
Lialoeh1129123-128,
Zar, J.H. (1974). Biostatistical analysis. Prentice-Hall Inc., Englewood Cliffs,
N.J. USA, 620 pp.
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MYCOTAXON
Volume XLVII, pp. 201-217 April-June 1993
HYPOGEOUS MYCORRHIZAL FUNGI OF SPAIN
Isabel F. Alvarez’, Javier Parladé', James M. Trappe’, and
Michael A. Castellano®
"Departamento de Patologia Vegetal. Institut de Recerca i Tecnologia Agroalimentaries
(IRTA). Centre de Cabrils. 08348 Cabrils, Barcelona, Spain.
7Department of Forest Science and Botany-Plant Pathology. Oregon State University, Corvallis,
Oregon 97331.
*United States Department of Agriculture, Forest Service, Pacific Northwest Research Station.
Forestry Sciences Laboratory, Corvallis, Oregon 97331.
ABSTRACT
Although scientific study of hypogeous fungi in Spain began early in this
century, our collections over five years (specially in the northern area of the
Iberian Peninsula) produced 18 new species records, including two new species
and one new genus. This paper includes first reports in Spain for E/aphomyces
muricatus, Glomus macrocarpum, Hymenogaster sp., Hymenogaster
albus, H. muticus, H. niveus, H. tener, Hysterangium clathroides, H.
gardneri, Loculotuber gennadi, Rhizopogon subareolatus, R.
ventricisporus, Tuber gennadii, T. levissimum, T. maculatum, T.
malencgonii and T. multimaculatum. Information on previously reported
species is expanded to other geographical areas. The updated relation of
Spanish hypogeous fungi and its ecological significance in the forests of northern
Spain are discussed.
KEY WORDS: Ascomycotina, Basidiomycotina, truffle, ectomycorrhiza,
Elaphomyces, Glomus, Hymenogaster, Hysterangium, Loculotuber,
Rhizopogon, Tuber.
202
INTRODUCTION
The study of hypogeous fungi in Spain was initiated by Lazaro Ibiza in
1908. The first published compilation of data available on hypogeous fungi in
Spain reported 34 species in 12 genera (Calonge et al. 1977). Subsequent
additions increased the number of reported fungi considerably (Calonge 1982,
Calonge et al. 1985b, Calonge 1990).
Most hypogeous fungi are desirable ectomycorrhizal symbionts of tree
seedlings because they increase water and nutrient uptake (Bowen 1973, Luoma
et al. 1991). Additional interest for their use in reforestation practices stems from
their capability to reduce soil drought stress in tree seedlings (Parke et al. 1983).
To use hypogeous fungi for increasing plantation success, they must be
collected in the field, identified, isolated or kept for spore inoculum, and
successfully infect seedlings in the nursery (Trappe 1977, Castellano et al. 1985,
Castellano and Molina 1989). Many hypogeous ectomycorrhizal fungi show plant-
fungus specificity (Molina and Trappe 1982).
Available information on hypogeous fungi in humid northern Spain has
been scarce. This area extends from Galicia to Catalonia, a total of ca. 14 million
ha. Presently, 4.4 million ha are covered with conifer, broadleaf and mixed
forests. An additional 3.3 million ha will be reforested in the future (Anonymous
1989).
The objective of this study was to collect and identify hypogeous
mycorrhizal fungi occurring with the major forest species of northern Spain for
possible later use in either conventional reforestation, or truffle production
programs.
MATERIALS AND METHODS
Fungi were collected at least twice a year from fall through spring of 1985-
1989. The Cantabric zone (from Galicia to the west part of the Pyrenees range)
was sampled from sea level to 1200 m elevation in a range of forest
associations. At the lower elevations (0 - ca 400 m) the stands consisted of either
pure Pinus pinaster Ait., Pinus radiata D. Don or a mixture of the two pines
with Castanea sativa Mill. At the middle elevations (300 - 700 m) similar mixed
forests were examined as also were plantations of Eucalyptus globulus Lab.
on the coastal slopes. Inland, in the Cantabric range, the tree species were Acer
pseudoplatanus L., Corylus avellana L., Fraxinus excelsior L., Ilex
aquifolium L., Populus spp., and Quercus petraea (Matts.) Liebl. At the
higher elevation (1000 - 1200 m) in Asturias (Picos de Europa) the forests were
composed of Fagus sy/vaticaL. and Quercus pyrenaica Willd. In Catalonia
(Montseny Range) the mixed forests were located at 1000-1200 m elevation and
composed of Abjes alba Mill, Fagus sylvatica L., Quercus ilex L.,
Quercus pubescens Willd. and plantations of Picea abies (L.) Karst. and
Pseudotsuga menziesii (Mirb.) Franco.
Other areas were occasionally searched for hypogeous fungi. Among them
203
were the Beceite range in southern Catalonia (1200 m elevation in a native
Pinus sylvestris L. forest) and plantations of introduced pines at the lower
elevations of the Montseny Range.
Hypogeous fungi were sought by raking the soil and litter. Care was taken
to replace the litter afterwards. Field data were taken on fresh sporocarp features,
overstory species composition, associated ectomycorrhizal hosts, and soil
characteristics. A few specimens were found with a trained dog. Some collections
provided by commercial truffle hunters lacked field data.
Because of their mycological interest, available collections of hypogeous
fungi from other areas of Spain recently described have been included in this
review (Alvarez et al. 1992). Hypogeous fungi from the Canary Islands were
compiled previously by Fogel (1980) and are not included here.
Microscopic observations were made by sectioning sporocarp tissue with
a razor blade and mounting the material in water, KOH, cotton blue or Melzer’s
reagent. Sporocarps were dried at 30°C for herbarium deposit.
Isolations were made from fresh sporocarps on MMN agar slants (Molina
and Palmer 1982). Herbarium specimens were deposited in the Instituto Antonio
José Cavanilles, Real Jardin Botanico (MA), Madrid, Spain and in the Oregon
State University Herbarium (OSC), Corvallis, Oregon. Other herbaria cited are the
National Fungus Collections, Washington D.C. (BPI), The Royal Botanic Gardens,
Kew (K), and the Istituto ed Orto Botanico, Torino (TO).
RESULTS
A total of 36 species of hypogeous fungi were collected and identified.
Eighteen are new records for Spain, including two new species.
The following list attempts to update the known species of hypogeous
fungi that occur in Spain. The list includes data on collection site characteristics
and associated mycorrhizal hosts when available, as well as the positive results
of isolation attempts.
Newly reported species and new localities (provinces) for previously
reported species are written in bold characters. Collection numbers, place of
deposit, and previous references are given in parentheses. An asterisk after the
locality indicates that it is an Autonomous Community (composed of one or more
provinces) and the exact locality is not known. The symbol (+) after the species
name indicates that it was isolated successfully. The identification of the species
reported by other authors has not been verified except when indicated.
Balsamia vulgaris Vitt.
BARCELONA, March 1988; 30-40 yr.- old C. avellana plantation on dry, stony pH 7.8 soil
(MA-28295, OSC), (Calonge et al. 1985b). CORDOBA, (Moreno et al. 1991).
Associated mycorrhizal hosts: Cistus albidus L. and C. avellana.
204
Choiromyces magnusii (Matt.) Paol.
EXTREMADURA%, (Calonge et al. 1985a).
Associated mycorrhizal hosts: unknown.
Chofiromyces venosus (Fries) Th. Fries (= C. meandriformis Vitt., Tuber
sinuosum Laz.)
ALAVA, (Calonge et al. 1977). BADAJOZ, (Lazaro Ibiza 1908, Moreno et al. 1991). CACERES,
(Moreno et al. 1991).
Associated mycorrhizal hosts: Cistus /ladanifer L., Fagus sp., and Quercus sp.
NOTE: The type collection of Tuber sinuosum Laz. is lost (Calonge et al. 1985a). Lazaro’s
illustrations and descriptions, however, are readily compatible with C. venosus.
Delastria rosea Tul.
BARCELONA, (Calonge et al. 1985b).
Associated mycorrhizal hosts: mixed forest, specific host unknown.
Elaphomyces anthracinus Vitt.
CORDOBA, (Moreno et al. 1991).
Associated mycorrhizal hosts: Q. i/ex.
Elaphomyces granulatus Fries
ASTURIAS, September 1986; in a Q. petraea and Populus sp. forest at 760 m. elevation,
soil pH:4 (MA-28299, OSC). GIRONA, (Unamuno 1941). PAIS VASCO%*, (Calonge et al. 1977).
PONTEVEDRA, November 1987; under P. pinaster on a controlled-burned, flat area. Soil
pH: 4.7 (MA-28297, MA-28298, OSC). SEGOVIA, (Calonge et al. 1977). SORIA, November
1987; under P. sy/vestris at 1200 m. elevation. Soil pH: 5.2 (MA-28296, OSC).
Associated mycorrhizal hosts: F. sylvatica, P. sylvestris, and Quercus sp.
Elaphomyces muricatus Fries
SANTANDER, May 1986; under Q. pyrenaica at 1200 m. elevation (MA-28300, OSC).
Associated mycorrhizal hosts: Q. pyrenaica.
Elaphomyces personii Vitt.
GIRONA, (Codina and Font-Quer 1931).
Associated mycorrhizal hosts: C. sativa.
Elaphomyces trappei Galan & Moreno
BADAJOZ, (Moreno et al. 1991).
Associated mycorrhizal hosts: C. /adanifer.
Gautieria mexicana (Fisch.) Zeller & Dodge
ALAVA, (Calonge et al. 1985b).
Associated mycorrhizal hosts: P. radiata.
Gautieria morchelliformis Vitt.
BARCELONA, (Aymerich and Llimona 1986). CORDOBA, (Moreno et al. 1991). CUENCA,
(Calonge et al. 1985a). LERIDA, (Calonge et al. 1977).
Associated mycorrhizal hosts: Abies sp., Pinus nigra Arnold, and Q. ilex.
Genea klotzschii Berk. & Broome
BARCELONA, February, March 1988; collected in a 30-40 year-old C. avel//ana plantation on
stony, pH 7.8 soil (MA-28301, MA-28302, OSC), (Calonge et al. 1985b). VALLADOLID,
(Calonge et al. 1985b).
Associated mycorrhizal hosts: C. ave//ana and Pinus pinea L.
205
Genea verrucosa Vitt.
CORDOBA, (Moreno et al. 1991). TOLEDO, (Arroyo et al. 1989).
Associated mycorrhizal hosts: C. albidus and Q. ilex.
Geopora arenicola (Lév.) Kers (= Sepultaria arenicola (Lév.) Cooke, S.
arenosa (Fuck.) Cooke)).
GIRONA, (Heim et al. 1934). GRANADA, (Moreno et al. 1986). MADRID, (Moreno et al. 1986).
MURCIA, (Honrubia et al. 1983). VALLADOLID, (Moreno et al. 1991).
Associated mycorrhizal hosts: Pinus spp., Pinus halepensis Mill. and P. pinaster.
Geopora cervina (Vel.) Schum.
GRANADA, (Moreno et al. 1986).
Associated mycorrhizal hosts: Quercus faginea Lamk. and Q. ilex.
Geopora cooperi Harkn.
ALAVA, (Mendaza and Diaz 1987). GRANADA, (Ortega et al. 1981). GUADALAJARA, (Moreno
et al. 1986). MADRID, (Moreno et al. 1991). MURCIA, (Honrubia et al. 1990).
Associated mycorrhizal hosts: Pinus spp. and Q. ilex.
Geopora foliacea (Schaeff.) Anmad (= Sepultaria foliacea (Schaeff.)
Boud.)
ALBACETE, (Honrubia et al. 1983). BARCELONA, (Heim et al. 1934). GRANADA, (Ortega et
al. 1981). GUADALAJARA, (Moreno et al. 1986). MADRID, (Moreno et al. 1986).
Associated mycorrhizal hosts: P. halepensis and Populus alba L.
Geopora nicaensis (Boud.) Torre
ALMERIA, (Calonge and Oria de Rueda 1988). BARCELONA, (Losa Quintana 1974, Moreno
et al. 1986). GRANADA, (Moreno et al. 1986). GUADALAJARA, (Moreno et al. 1991). MADRID,
(Moreno et al. 1986).
Associated mycorrhizal hosts: conifers and Q. i/ex.
Geopora sumneriana (Cook.) Torre
GRANADA, (Moreno et al. 1986). MADRID, (Moreno et al. 1986).
Associated mycorrhizal hosts: Cedrus sp.
Geopora tenuis (Fuck.) Gal.& Ort.
GRANADA, (Galan and Ortega 1982).
Associated mycorrhizal hosts: unknown.
Glomus macrocarpum Tul. & Tul.
ASTURIAS, September 1986; in a mixed forest of A. pseudoplatanus, F. excelsior, and
Q. petraea at 700 m. elevation. Soil pH: 4 (MA-28303, OSC).
Associated mycorrhizal hosts: unknown.
Glomus microcarpum Tul. & Tul. (= Endogone microcarpa (Tul. & Tul.)
Tul.& Tul.
BARCELONA, (Heim et al. 1934).
Associated mycorrhizal hosts: unknown.
Hydnangium aurantiacum Heim & Mal., (= H. carotaecolor Berk.)
BARCELONA, (Codina and Font-Quer 1931, Heim et al. 1934).
Associated mycorrhizal hosts: P. halepensis and Q. ilex.
206
Hydnangium carneum Wallr. 4
BADAJOZ, (Moreno et al. 1991). LA CORUNA, (Jeppson 1987).
Associated mycorrhizal hosts: Eucalyptus spp.
Hydnangium pila Pat.
GIRONA, (Heim et al. 1934).
Associated mycorrhizal hosts: Quercus suber L.
NOTE: The generic disposition of this species is disputable by modern concepts. Svrcek’s
designation of it as Octaviania pila (Pat.) Svrc. is nomenclaturally unacceptable by
International Code of Botanical Nomenclature (Svrcek 1958).
Hydnotrya tulasnei (Berk.) Berk. & Broome
BARCELONA, (Calonge et al. 1985a). VALLADOLID, (Calonge 1982). VIZCAYA, (Mendaza and
Diaz 1987).
Associated mycorrhizal hosts: C. avellana, F. sylvatica, Pinus spp. and Quercus spp.
Hymenogaster sp. No. 1 :
GIRONA, November, December 1989; under P. menziesii at 1100 m. elevation in pH 5.5 soil
(MA-28314, MA-28315, MA-28316, MA-28317, OSC). PONTEVEDRA, November 1987; in a
15-yr.-old plantation of P. menziesii on flat ground with no understory (MA-28313, OSC).
Associated mycorrhizal hosts: P. menziesii.
Hymenogaster sp. No. 2
GIRONA, October 1989; under P. menziesii at 1100 m. elevation in pH 5.5 soil (MA-28318,
OSC).
Associated mycorrhizal hosts: P. menziesii.
Hymenogaster aibus Berk. & Br.
ASTURIAS, October 1986; under E. globulus at 500 m. elevation (MA-28304, OSC).
Associated mycorrhizal hosts: E. globulus.
Hymenogaster bulliardi Vitt.
TOLEDO, (Calonge et al. 1985b).
Associated mycorrhizal hosts: unknown.
Hymenogaster calosporus Tul. & Tul.
BARCELONA, (Heim et al. 1934).
Associated mycorrhizal hosts: C. avellana, Q. ilex and Sorbus aucuparia L.
Hymenogaster cerebellum Cavara
CACERES, (Moreno et al. 1991).
Associated mycorrhizal hosts: Q. pyrenaica.
Hymenogaster citrinus Vitt.
ASTURIAS, May 1986; under Quercus robur L. at 160 m elevation (MA-28306, OSC).
BARCELONA, (Heim et al. 1934). CORDOBA, (Moreno et al. 1991). GIRONA, October 1987;
under F. sy/vatica at 1100 m elevation (MA-28305, OSC).
Associated mycorrhizal hosts: F. sylvatica, Q. ilex and Q. robur.
Hymenogaster muticus Berk. & Br.
GIRONA, December 1987; under P. sy/vestris (MA-28307, OSC).
Associated mycorrhizal hosts: P. sy/vestris.
207
Hymenogaster niveus Vitt.
SANTANDER, May 1986; under F. sy/vatica at 1200 m elevation (MA-28308, OSC).
Associated mycorrhizal hosts: F. sylvatica.
Hymenogaster olivaceus Vitt.
BARCELONA, (Heim et al. 1934).
Associated mycorrhizal hosts: C. avellana and Q. ilex.
Hymenogaster populetorum Tul. & Tul.
BARCELONA, (Calonge et al. 1985b).
Associated mycorrhizal hosts: unknown.
Hymenogaster remyi Zeller & Dodge in Dodge & Zeller
GUADALAJARA, (Calonge 1982).
Associated mycorrhizal hosts: Pinus sp.
Hymenogaster tener Berk. & Br.
ASTURIAS, May 1986; under mixed forest of E. giobulus and P. radiata at475m elevation,
October 1986; under F. sy/vatica at 600 m. elevation (MA-28309, MA-28310, OSC).
Associated mycorrhizal hosts: F. sylvatica.
Hymenogaster vulgaris Tul.& Tul. in Berk. & Br.
BARCELONA, (Heim et al. 1934, Calonge et al. 1985a).
GIRONA, November, December 1989; under P. menziesii at 1100 m. elevation in pH 5.5 soil
(MA-28311, MA-28312, OSC).
Associated mycorrhizal hosts: Pinus sp. and P. menziesii.
Hysterangium clathroides Vitt.
ASTURIAS, May 1986; in a 15-years-old P. radiata plantation, at 270 m. elevation. (MA-
28319, OSC).
Associated mycorrhizal hosts: P. radiata.
Hysterangium gardneri Fischer
ASTURIAS, May 1986; in a mixed stand of C. sativa and E. globulus at 575 m. elevation
(MA-28320, OSC).
Associated mycorrhizal hosts: E. globulus.
Hysterangium rickenii Soehn.
GRANADA, (Moreno et al. 1986).
Associated mycorrhizal hosts: Q. faginea and Q. ilex.
Hysterangium separabile Zeller
CUENCA, (Calonge et al. 1985a). MADRID, (Calonge et al. 1977).
Associated mycorrhizal hosts: P. nigra and P. sylvestris.
Hysterangium stoloniferum Tul. var. rubescens (Quél.) Zeller & Dodge.
CACERES, (Moreno et al. 1991). CORDOBA, (Moreno et al. 1991).
Associated mycorrhizal hosts: C. sativa and Q. ilex.
Hysterangium thwaitesii Berk. & Broome
ALAVA, (Calonge et al. 1977).
Associated mycorrhizal hosts: unknown.
208
Labyrinthomyces donkii Mal.
ALICANTE, (Honrubia 1984). CADIZ, (Calonge 1982, 1983). MURCIA, (Honrubia 1984).
Associated mycorrhizal hosts: E. g/obulus.
NOTE: This genus has been recently revised by Trappe et al. (1992).
Leucogaster floccosus Hesse
MURCIA, (Honrubia and Llimona 1981).
Associated mycorrhizal hosts: unknown.
Loculotuber gennadi Trappe, Parladé & Alvarez
CACERES, May 1987; (MA-26771, OSC). (Alvarez et al. 1992).
Associated mycorrhizal hosts: unknown.
Martellia mediterranea Moreno, Galan & Montecchi
CACERES, (Moreno et al. 1991).
Associated mycorrhizal hosts: C. /adanifer and Q. ilex.
Melanogaster ambiguus (Vitt.) Tul. (+).
BARCELONA, (Calonge 1982). GIRONA, November 1989 and June 1990 under P. menziesii
at 1100 m. elevation in pH 5 soil (MA-28321, MA-28322, MA-28323, MA-28324, MA-28325,
OSC), (Martin 1988). MADRID, (Moreno et al. 1986).
Associated mycorrhizal hosts: F. sy/vatica, P. menziesii, Q. ilex, and Q. suber.
Melanogaster broomeianus Berk. in Tul. & Tul
ALBACETE, (Calonge and Demoulin 1975). BARCELONA, (Calonge et al. 1985b, Llistosella
and Aguasca 1986). GIRONA, (Maire 1937, Singer 1947). MADRID, (Calonge et al. 1985a).
Associated mycorrhizal hosts: P. sy/vestris and P. pinea.
Melanogaster variegatus (Vitt.) Tul. & Tul.
ALBACETE, (Calonge 1973). CACERES, (Moreno et al. 1991). CORDOBA, (Moreno et al.
1991). GUIPUZCOA, (Lazaro Ibiza 1912). MURCIA, (Honrubia et al. 1990). VALENCIA, (Lazaro
Ibiza 1912).
Associated mycorrhizal hosts: Pinus spp. and Q. suber.
Octaviania asterosperma Vitt.
CACERES, (Moreno et al. 1991).
Picoa juniperi Vitt.
MADRID, (Calonge et al. 1985a). ZARAGOZA, (Calonge 1982).
Associated mycorrhizal hosts: unknown.
Rhizopogon /uteolus Fries & Nordh. (+)
ALAVA, December 1989; under P. sy/vestris (MA-28326, OSC). ASTURIAS, October 1986:
under P. radiata at 100 m. elevation (MA-28327). BARCELONA, (Maire et al 1933, Heim et
al. 1934, Llimona 1984, Martin 1988). GIRONA, (Maire et al. 1933, Llimona 1983, Martin
1988). LERIDA, (Maire 1937). MADRID, (Calonge and Zugaza 1973, Calonge and Demoulin
1975). PONTEVEDRA, November 1985, 1986, and 1987; under P. pinaster in compacted,
sandy pH 4.7-6 soils and sometimes abundant in controlled-burned areas (MA-28328 MA-
28329 MA-28330, MA-28331, MA-28332, MA-28333, MA-28334, MA-28335, MA-28336, OSC).
TARRAGONA, (Maire 1937).
Associated mycorrhizal hosts: P. nigra, P. pinaster, P. pinea, P. radiata and P.
sylvestris.
209
Rhizopogon roseolus (Corda ex Sturm) Th. Fries (= R. aestivus (Tul. &
Tul.) Tul. & Tul., R. rubescens Tul & Tul.) (+)
ALBACETE, (Honrubia et al. 1982). ALICANTE, (Honrubia et al. 1982). ALMERIA, (Honrubia
et al. 1982). ASTURIAS, December 1985, May and October 1986; under P. radiata
plantations at 100-350 m. elevation in pH 5.1 soil (MA-28337 and MA-28338, OSC).
BALEARES, (Malengon and Bertault 1972). BARCELONA, (Codina and Font-Quer 1931, Maire
et al. 1933, Heim et al. 1934, Maublanc 1936, Singer 1947, Calonge and Demoulin 1975,
Tabares and Rocabruna 1987, Martin 1988). GIRONA, (Maire et al. 1933, Maire 1937, Font-
Quer 1954, Calonge and Demoulin 1975, Llimona 1983, Martin 1988). JAEN, (Honrubia et al.
1982). LEON, November 1989; under P. sy/vestris in sandy soil (MA-28342, MA-28343,
OSC). LERIDA, (Calonge and Demoulin 1975, Llimona 1983, Martin 1988). MADRID, (Calonge
1973, Calonge and Zugaza 1973, Calonge and Demoulin 1975). MALAGA, (Calonge and
Demoulin 1975). MURCIA, (Honrubia et al. 1982). PONTEVEDRA, November 1986; under P.
pinaster (MA-28339, OSC). TARRAGONA, September 1987, August 1989; under P.
sylvestris at 1100 m. elevation in pH 7.4 soil (MA-28340, MA-28341, OSC). VALLADOLID,
(Calonge and Demoulin 1975).
Associated mycorrhizal hosts: P. halepensis, Pinus mugo Turra., P. nigra, P. pinaster,
P. pinea, P. radiata and P. sylvestris.
Rhizopogon subareolatus Smith (+)
GIRONA, November, December 1987 and 1988, abundant under P. menziesii plantations
or under Abies sp. with P. menziesii present on slopes at 1100 m. altitude in pH 4.7-6 soil
(MA-28344, MA-28345, MA-28346, MA-28347, MA-28348, MA-28349, MA-28350, and MA-
28351, OSC).
Associated mycorrhizal hosts: P. menziesii.
NOTE: This species has not previously been reported in continental Europe. It was reported
in the Madeira islands where it was probably introduced with P. menziesii (Fogel 1980).
Rhizopogon ventricisporus Smith (+)
TARRAGONA, September 1987; under P. sy/vestris at 1100 m. elevation in pH 7.4 soil (MA-
28352, OSC).
Associated mycorrhizal hosts: P. sylvestris.
Rhizopogon vulgaris (Vitt.) M. Lange (= R. provincialis Tul. & Tul.) (+)
ALBACETE, (Honrubia et al. 1982). ASTURIAS, May 1986 and October 1986; under P.
radiata at 100-350 m elevation in pH 5.1 soil (MA-28353, MA-28354, MA-28355 and MA-
28356, OSC). BARCELONA, (Codina and Font-Quer 1931, Martin 1988). GIRONA, (Heim et
al. 1934, Malencon and Bertault 1971). MADRID, (Calonge and Demoulin 1975). MALAGA,
(Calonge and Demoulin 1975). PONTEVEDRA, November 1985, 1986 and 1987; under P.
pinaster on coast in pH 4.7 soil in controlled-burned areas and under P. menziesii in 15
year-old plantations (MA-28357, MA-28358, MA-28359, MA-28360 and MA-28361, OSC).
SORIA, November 1987; under P. sy/vestris at 1200 m elevation in pH 5.2 soil (MA-28362
and MA-28363, OSC).
Associated mycorrhizal hosts: P. halepensis, P. nigra, P. pinaster, P. radiata, P.
sylvestris and P. menziesii.
Terfezia arenaria (Mor.) Trappe (= T. /eonis Tul.)
BADAJOZ, (Calonge et al. 1977, Calonge et al. 1985a). CACERES, May 1987 (MA-28364 and
MA-28365, OSC), (Moreno et al. 1986). HUELVA, (Calonge et al. 1985a). TOLEDO, (Calonge
et al. 1985b).
Associated mycorrhizal hosts: C. Jadanifer, Cistus monspeliensisL., Cistus salvifolius
L. and Helianthemum guttatum Mill.
210
Terfezia berberiodora (Lesp. ex Tul. & Tul.) Tul. & Tul.
LA CORUNA, (Castro and Freire 1982).
Associated mycorrhizal hosts: unknown.
Terfezia claveryi Chat.
BURGOS, (Calonge et al. 1985b). GRANADA, (Calonge et al. 1985b, Moreno et al. 1986).
MADRID, (Calonge et al. 1985b).
Associated mycorrhizal hosts: Helianthemum sSsalicifolium Pers.
Terfezia hispanica Laz.
ALBACETE, (Lazaro Ibiza 1908). BADAJOZ, (Lazaro Ibiza 1908). CACERES, (Lazaro Ibiza
1908). CIUDAD REAL, (Lazaro Ibiza 1908). CORDOBA, (Lazaro Ibiza 1908). GRANADA,
(Lazaro Ibiza 1908). GUADALAJARA, (Lazaro Ibiza 1908). MADRID, (Lazaro Ibiza 1908).
SALAMANCA, (Lazaro Ibiza 1908). TOLEDO, (Lazaro Ibiza 1908). ZARAGOZA, (Lazaro Ibiza
1908).
Associated mycorrhizal hosts: Helianthemum paniculatum Durm., H. salicifolium,
Helianthemum vulgaris Gaertn., and Tuberaria variabilis Wk.
NOTE: T. hispanica may be an aberrant collection of 7. o/biensis; the type collection is lost
(Calonge et al. 1985a). Because of its distinctive vinaceous color as illustrated by Lazaro Ibiza
(1908), however, we prefer to keep it separate from 7. o/biensis pending new collections.
Terfezia olbiensis (Tul. & Tul.) Tul. & Tul. (= T. feptoderma (Tul. & Tul.)
Tul. & Tul., 7. cadevalli Font-Quer, Tuber lutescens Laz., T. pallidumLaz.)
ALAVA, (Calonge et al. 1977). ALBACETE, (Lazaro Ibiza 1908). AVILA, (Lazaro Ibiza 1908).
BADAJOZ, (Calonge et a!. 1977, Calonge et al. 1985b). BARCELONA, (Codina and Font-Quer
1931). BURGOS, (Calonge et al. 1985b). CACERES, May 1987 (MA-28366 and MA-28367,
OSC), (Moreno et al. 1986). CIUDAD REAL, (Lazaro Ibiza 1908). MADRID, (Lazaro Ibiza 1908,
Calonge 1982, Calonge et al. 1985b). SALAMANCA, (Lazaro !biza 1908). SEGOVIA, (Calonge
et al. 1985b). TOLEDO, (Calonge et al. 1985b). VALLADOLID, (Lazaro Ibiza 1908). ZAMORA,
(Lazaro ibiza 1908).
Associated mycorrhizal hosts: H. guttatum, H. paniculatum, Q. ilex, and T. variabilis.
NOTE : We have found many sporocarps intermediate in morphology between those reported
for 7. olbiensis and T. leptoderma in the numerous collections we have examined. Most
differences appear to be a matter of developmental stage, as indicated by Fogel (1980).
Hence, we regard 7. /eptoderma as a synonym of 7. olbiensis (we choose T. o/biensis
as the epithet to retain, because its type collection contains more mature material than does
that of 7. /Jeptoderma). Although the type collections of Terfezia cadevalli Font-Quer,
Tuber lutescens Laz. and T. pallidum Laz. are lost (Calonge et al., 1985a), the descriptions
of each present no reason to regard them other than as somewhat inmature to mature stages
of 7. olbiensis.
Trappea darkeri (Zeller) Castellano
GUADALAJARA, (Calonge 1982).
Associated mycorrhizal hosts: Pinus sp.
NOTE: This collection was initially reported as Hysterangium membranaceum Vitt.
(Castellano 1990).
Tuber aestivum Vitt.
BARCELONA, March 1987 (MA-28368, OSC), (Unamuno 1941, Calonge et al. 1985a).
CORDOBA, (Moreno et al. 1991). GIRONA, (Codina and Font-Quer 1931, Unamuno 1941).
LERIDA, March 1987; under Pinus sp. (MA-28369, OSC). TERUEL, March 1987; under
Pinus sp. (MA-28370, OSC).
Associated mycorrhizal hosts: C. avellana, Pinus sp. and Q. ilex.
211
Tuber asa Tul. & Tul.
BADAJOZ, (Moreno et al. 1991). CACERES, May 1987, (MA-28371 and MA-28372, OSC),
(Moreno et al. 1991).
Associated mycorrhizal hosts: C. ladanifer.
Tuber borchii Vitt.
MURCIA, (Honrubia and Llimona 1981).
Associated mycorrhizal hosts: P. halepensis.
Tuber brumale Vitt.
BARCELONA, March 1988; under C. avellana in 30-40 year-old plantations in pH 7.8 soil
(MA-28373, OSC), (Codina and Font-Quer 1931), and three collections at TO. CASTELLON,
(Calonge et al. 1977). GIRONA, (Lazaro Ibiza 1912, Codina and Font-Quer 1931, Heim et al.
1934, Unamuno 1941). GUADALAJARA, (Calonge et al. 1977). MADRID, one collection at TO.
TARRAGONA, March 1987 (MA-28374, OSC).
Associated mycorrhizal hosts: C. ave/l/ana and Quercus sp.
Tuber excavatum Vitt.
BARCELONA, (Calonge et al. 1985a, b). CORDOBA, (Moreno et al. 1991). GUADALAJARA,
December 1987, under Populus sp. at 1100 m. altitude (MA-28375, OSC).
Associated mycorrhizal hosts: C. avellana, Pinus sp., Populus sp. and Q. ilex.
Tuber foetidum Vitt.
BALEARES, (Calonge 1982, 1983). BARCELONA, (Calonge et al. 1985b).
Associated mycorrhizal hosts: Pinus sp.
Tuber gennadii (Chat.) Pat.
CACERES, May 1987 (MA-28376, OSC).
Associated mycorrhizal hosts: unknown.
Tuber levissimum Gilkey
GIRONA, November 1989, December 1989, and June 1990; under P. menziesii in pH 5.5
soil (MA-28377, MA-28378, MA-28379, MA-28380, OSC).
Associated mycorrhizal hosts: P. menziesii.
Tuber maculatum Vitt.
GIRONA, October, December 1989; under P. menziesii in pH 5.5 soil (MA-28381, MA-28382,
OSC).
Associated mycorrhizal hosts: P. menziesii.
Tuber malenconi Don., Riouss. & Chev.
BARCELONA, March 1987 (MA-28385, OSC). LERIDA, March 1987 (MA-28384, OSC).
TERUEL, March 1987 (MA-28383, OSC).
Associated mycorrhizal hosts: unknown.
Tuber maresa Font-Quer
GIRONA, (Codina and Font-Quer 1931).
Associated mycorrhizal hosts: unknown.
NOTE: 7. maresa seems likely to be an immature collection of 7. mesentericum, but we
have not located the holotype for study.
Tuber melanosporum Vitt.
BARCELONA, (Unamuno 1941), one collection at BP! (Lloyd 22118) pius five collections at
TO. GIRONA, (Lazaro Ibiza 1912, Codina and Font-Quer 1931, Heim et al. 1934).
212
GUADALAJARA, December 1987, under Q. i/ex at 1200 m. altitude (MA-28390, OSC),
(Calonge et al. 1985b, Moreno et al. 1986). HUESCA, December 1987; under Q. ilex
(MA-28386, MA-28387, MA-28388, MA-28391, OSC). TARRAGONA, February - March 1987;
under C. sativa (MA-28392 and MA-28393, OSC). TERUEL, March, 1987 (MA-28389, OSC).
Associated mycorrhizal hosts: C. sativa, C. avellana, and Q. ilex.
Tuber mesentericum Vitt.
BARCELONA, (Unamuno 1941). TARRAGONA, March 1987; under C. sativa (MA-28394,
OSC). TERUEL, (Calonge 1983).
Associated mycorrhizal hosts: C. sativa.
Tuber multimaculatum Parladé, Trappe & Alvarez
TARRAGONA, March 1987; (MA-26769, MA-26770, OSC). (Alvarez et al. 1992).
Associated mycorrhizal hosts: unknown.
Tuber nitidum Vitt.
CORDOBA, (Moreno et al. 1991).
Associated mycorrhizal hosts: Q. i/ex.
Tuber oligospermum (Tul. & Tul.) Trappe (= Terfezia oligosperma Tul.
& Tul.)
LA CORUNA, (Castro and Freire 1982).
Associated mycorrhizal hosts: unknown.
Tuber panniferum Tul. & Tul
CORDOBA, (Moreno et al. 1991).
Associated mycorrhizal hosts: Q. i/ex.
Tuber puberulum Berk. & Broome
BARCELONA, (Heim et al. 1934).
Associated mycorrhizal hosts: Q. lex.
Tuber rufum Pico:Fries (= Tuber ferrugineum Vitt.)
BARCELONA, March 1988; in a 30-40 yr.-old C. ave//ana plantation in a stony pH 7.8 soil
(MA-28396, OSC), (Calonge et al. 1985a, b; Moreno et al. 1991). CACERES, (Moreno et al.
1991). CASTELLON, (Calonge et al. 1977). CUENCA, (Calonge et al. 1977). GIRONA,
November, December 1989; under P. menziesii in Ph 5.5 soil (MA-28395, MA-28397, OSC).
MALAGA, one collection at K.
Associated mycorrhizal hosts: C. sativa, C. avellana, Pinus sp., Q. faginea, and Q. ilex.
DISCUSSION
Throughout five years of field sampling in forests of northern Spain, we
observed that hypogeous fungi are scarce and difficult to find. Only species of
the genus Rhizopogon were obtained with relative ease every fall and spring,
once the field sites where they occur had been identified. The finds of two new
taxa of hypogeous Pezizales (Tuberales) and new records of others indicate that
systematic research is needed in many areas of Spain to complete the list of
hypogeous mycoflora.
Animal digging (holes, channels or scraped soil) was observed in mild
215
coastal areas of Asturias and Galicia and in mountains of Catalonia. Animal
activity of this sort was not observed at other places where hypogeous fungi
were collected. The interrelationship between mycophagous small mammals and
hypogeous fungi has been proven in North America (Fogel and Trappe 1978,
Maser et al. 1978, Ure and Maser 1982) and in Australia (Malajczuk et al. 1987).
Also, wild boars and foxes readily penetrate commercial plantations of Tuber
melanosporum protected with electric fences in search of sporocarps. The
relationship between animal vectors and hypogeous fungi has not been studied
in Spain, although Durrieu et al. (1984) report substantial utilization by small
mammals in the French Pyrenees. The scarcity of sporocarps observed in many
Spanish forests may be because of the low number of species, or low
populations of animal vectors thus reducing spore dissemination of fungi already
present in the forests.
Many genera of hypogeous fungi, both Ascomycetes and Basidiomycetes,
have never been found in Spain. This may indicate lack of long term studies on
the hypogeous mycoflora of Spanish forests. Hunt and Trappe (1987) indicate
that documenting all species of hypogeous ectomycorrhizal fungi in a forest stand
requires long-term collecting over several years. The production of sporocarps
varies from year to year (Fogel 1976) and hypogeous fungal species are scarce
when compared to epigeous species in stands of conifers (Luoma et al. 1991).
The relatively low diversity of tree species (particularly conifers) in Spanish forests
compared with that in places such as the American Northwest, where hypogeous
fungi abound, could also explain in part the low species diversity of hypogeous
— fungi in Spain.
Northern Spain has residual virgin forests, but most extensive forests are
the result of human reforestation, often with species native to other continents
such as Pinus radiata, P. lambertiana, Pseudotsuga menziesii, and
Eucalyptus spp. Research programs to introduce exotic forest species into
Europe are likely to continue, given the poverty of indigenous genetic material
available (CEE 1984). Hence, parallel research on the hypogeous mycorrhizal
symbionts of forest tree species in their area of origin and on the native
hypogeous fungi in the introduction area is desirable to assess the need of
introducing a particular fungal symbiont for successful establishment of the trees.
In some Spanish plantations of P. menziesii, the host-specific hypogeous
mycorrhizal fungi Rhizopogon subareolatus has been collected consistently
and was probably introduced with its host seedlings long ago.
Isolations from sporocarps were obtained with relative ease only from the
Rhizopogon species. Some species such as R. roseo/us, were collected in
both acid (pH 5) and slightly basic (pH 7.4) soils. This genetic diversity is an
important consideration in selecting fungal candidates for use in nursery
inoculations (Trappe 1977). Knowledge of the adaptability and host range of the
fungal symbiont as well as the receptivity and infectivity of the planting site are
important in evaluating the success of introduced fungi in improving forestry
practices.
214
ACKNOWLEDGEMENTS
Financial support for this research was provided by U.S.- Spain Joint Committee
for Scientific and Technological Cooperation grant No. CCA-84 11/013. We thank
Drs. R. Fogel and C. Walker for their constructive comments on the manuscript.
LITERATURE CITED
Alvarez, |.F., J. Parladé and J.M. Trappe. 1992. Locu/otuber gennadii gen.
& comb. nov. and Tuber multimaculatum sp. nov. (Tuberaceae).
Mycologia 84:926-929.
Anonymous. 1989. Anuario de Estadistica Agraria. Minist. Agric. Pesca
Aliment. Madrid. 678 pp.
Arroyo, |., F.D. Calonge, G. Carrascosa and E. Sau. 1989. Nuovi funghi ipogei
di Spagna Ill. Hydnocystis clausa (Tul.) Cerutie Genea verrucosa Vitt.
Micol. Ital. 1:3-7.
Aymerich, J. and X. Llimona. 1986. Gautieria morchellaeformis Vitt.
(Gastrosporiales, Gasteromicetes) al Bergueda (Catalunya). Folia Bot. Misc.
5:145.
Bowen, G.D. 1973. Mineral nutrition of ectomycorrhizae. In Ectomycorhizae.
Their ecology and physiology. Edited by G.C. Marks and T.T. Kozlowski.
Academic Press, New York. pp. 151-205.
Calonge, F.D. 1973. Estudios sobre hongos. IV. Aportacion al catalogo de las
provincias de Madrid y Sevilla. An. Inst. Bot. Cavanilles 30: 19-32.
Calonge, F.D. 1982. Adiciones al catalogo de hongos hipogeos de Espana.
Garcia de Orta Ser. Estud. Agron. 9:143-146.
Calonge, F.D. 1983. Algunos hongos raros 0 nuevos para Espana. Lazaroa
5:283-289.
Calonge, F.D. 1990. Check-list of the Spanish Gasteromycetes (Fungi,
Basidiomycotina). Crypt. Bot. 2:33-55.
Calonge, F.D. and V. Demoulin. 1975. Les Gasteromycetes d’Espagne. Bull.
Soc. Mycol. Fr. 91:247-292.
Calonge, F.D. and J.A. Oria de Rueda. 1988. Aportacion a la micoflora de la
provincia de Almeria. Bol. Soc. Micol. Cast. 12:93-106.
Calonge, F.D. and A. Zugaza. 1973. Catalogo de los hongos presentados en la
| exposicion de Madrid, del 1 al 5 de Noviembre de 1972. An. Inst. Bot.
Cavanilles 30:33-55.
Calonge, F.D., A. Rocabruna and M. Tabarés. 1985a. Nuevos datos sobre los
hongos hipogeos de Espana. Bol. Soc. Micol. Cast. 9:45-54.
Calonge, F.D., A. Rocabruna, M. Tabarés and N.B. Rodriguez. 1985b. Nuevos
datos sobre los hongos hipogeos de Espana. Il. Géneros Balsamia,
Delastriay Genea, novedades para el catalogo espanol. Butll. Soc. Cat.
Micol. 9:57-64.
Calonge, F.D., M. de la Torre and M. Lawrynowicz. 1977. Contribucién al estudio
de los hongos hipogeos de Espana. An. Inst. Bot. Cavanilles 34:15-31.
215
Castellano, M.A. 1990. The new genus Trappea_ (Basidiomycotina,
Hysterangiaceae), a segregate from Hysterangium. Mycotaxon 38: 1-9.
Castellano, M.A. and R. Molina. 1989. Mycorrhizae. In: The container tree
nursery manual, Vol. 5, Handbk. 674. Edited by T.D. Landis, R.W. Tinus,
S.E. McDonald and J.P. Barrett. U.S.D.A. For. Serv., Washington, D.C.
pp. 101-167.
Castellano, M.A., J.M. Trappe and R. Molina. 1985. Inoculation of container-
grown Douglas-fir with basidiospores of Rhizopogon vinicolor and R.
colossus: effects of fertility and spore application rate. Can. J. For. Res.
15:10-13.
Castro, M.L. and L. Freire. 1982. Terfezia berberiodora y T. oligosperma
encontradas en Galicia, nuevas para Espana. Collect. Bot. (Barc.) 13:44 1-
444.
CEE. 1984. L’Europe verte. La Couverture verte de L’Europe. Nos fdrets
d’aujourd’hui et de demain. Brussels. 24 pp.
Codina, J. and P. Font-Quer. 1931. Introduccié a l’estudi dels macromicets de
Catalunya. Cavanillesia 3: 100-189.
Durrieu, G., M. Genard and F. Lescourret. 1984. Les micromammiféres et la
symbiose mycorrhizienne dans une forét de montagne. Bul. Ecol. 15:253-
263.
Fogel, R. 1976. Ecological studies of hypogeous fungi. Il. Sporocarp phenology
in a western Oregon Douglas-fir stand. Can. J. Bot. 54:1152-1162.
Fogel, R. 1980. Additions to the hypogeous mycoflora of the Canary Islands and
| Madeira. Contr. Univ. Mich. Herb. 14:75-82.
Fogel, R. and J.M. Trappe. 1978. Fungus consumption (mycophagy) by small
mammals. Northwest Sci. 52:1-31.
Font-Quer, P. 1954. Plantas de Greixa (Pirineo catalan). Collect. Bot. (Barc.)
4:173-177.
Galan, R. and A. Ortega. 1982. Geopora tenuis (Fuckel) Galan & Ortega
comb. nov., nuevo pezizal para la micoflora espanola. Bol. Soc. Micol.
Cast. 7:49-51.
Heim, R., P. Font-Quer and J. Codina. 1934. Fungi Iberici. Observations sur la
flore mycologique catalane. Treb. Mus. Cienc. Nat. Barc, Ser. Bot. 15,
146 pp.
Honrubia, M. 1984. Labyrinthomyces donkii Malengon, en el S.E. de
Espana. Int. J. Mycol. Lichenol. 1:345-349.
Honrubia, M. and X. Llimona. 1981. Aportacion al conocimiento de los hongos
del S.E. de Espafia. IV. Tres citas nuevas para la microflora espafola.
Pustularia insignis, Tuber borchii, Leucogaster cf. floccosus. An.
Univ. Murcia Cienc. 37:81-90.
Honrubia, M., P. Berthet and X. Llimona. 1983. Contribution a la connaissance
des champignons du sud-est de I’Espagne. VII. Pezizales (Ascomycetes).
Bull. Soc. Linn. Lyon 52:46-62.
Honrubia, M., F.D. Calonge, V. Demoulin, G. Moreno and X. Llimona. 1982.
Aportacion al conocimiento de los hongos del S.E. de Espana VI:
Esclerodermatales, Licoperdales, Nidulariales, Falales, Himenogasterales,
Podaxales (Gasteromicetes, Basidiomicetes). An. Univ. Murcia Cienc.
38: 101-132.
216
Honrubia, M., A. Cano and C. Molina-Nifirola. 1990. Nota sobre hongos
hipogeos del semiarido murciano. Resumenes V Reunién Conjunta de
Micologia. Rev. Iberoamericana Micol. 7:95.
Hunt, G.A. and J.M. Trappe. 1987. Seasonal hypogeous sporocarp production
in a western Oregon Douglas-fir stand. Can. J. Bot. 65:438-445.
Jeppson, M. 1987. Notes on some Spanish Gasteromycetes. Bol. Soc. Micol.
Madr. 11:267-282.
Lazaro Ibiza, B. 1908. Nuevos Tuberaceos de Espana. Rev. R. Acad. Cienc.
Exactas Fis. Nat. Madr. 6:801-826.
Lazaro Ibiza, B. 1912. Notas micoldgicas. Coleccién de datos referentes a los
hongos de Espafia. 3° serie. Mem. R. Soc. Esp. Hist. Nat. 7:287-341.
Llimona, X. 1983. Sobre fongs de primavera a Catalunya. Butll. Soc. Cat. Micol.
7:33-45.
Llimona, X. 1984. Llista d’espécies de l’exposicio de fongs organitzada a
Barcelona l’octubre de 1977. Butll. Soc. Cat. Micol. 8:39-46. ,
Llistosella, J. and M. Aguasca. 1986. El 1° mini foray de la British Mycological
Society a Catalunya. Butll. Soc. Cat. Micol. 10:19-33.
Losa Quintana, J.M. 1974. Contribuci6n al conocimiento de los Ascomycetes con
apotecios de Catalufia. Collect. Bot. (Barc.) 9:45-60.
Luoma, D.L., R.E. Frenkel, and J.M. Trappe. 1991. Fruiting of hypogeous fungi
in Oregon Douglas-fir forests: seasonal and habitat variation. Mycologia 83:
335-353.
Maire, R. 1937. Fungi Catalaunici. Series Altera. Contribution a l'étude de la Flore
Mycologique de la Catalogne. Publ. Inst. Bot. Barc. 3, 128 pp.
Maire, R., J. Codina and P. Font-Quer. 1933. Fungi Catalaunici. Serie Botanica.
Contributions a l’étude de la Flore Mycologique de la Catalogne. Publ. Mus.
Cienc. Nat. Barc. 15: 1-120.
Malajczuk, N., J.M. Trappe and R. Molina. 1987. Interrelationships among some
ectomycorrhizal trees, hypogeous fungi and small mammals. Western
Australian and northwestern American parallels. Aust. J. Ecol. 12:53-55.
Malengon, G. and R. Bertault. 1971. Champignons de la Peninsule Ibérique. I.
Explorations entre le Midi valencien et le Montseny. Acta Phytotaxon. Barc.
8: 1-68.
Malencon, G. and R. Bertault. 1972. Champignons de la Peninsule ibérique. IV.
Les lles Baléares. Acta Phytotaxon. Barc. 11:1-64.
Martin, M.P. 1988. Aportacidn al conocimiento de las higroforaceas y los
gasteromicetos de Catalufia. Edic. Esp. Soc. Cat. Micol. 2. 508 pp.
Maser, C., J.M. Trappe and R.A. Nussbaum. 1978. Fungal-small mammal
interrelationships with emphasis on Oregon coniferous forests. Ecology
59:799-809.
Maublanc, M.A. 1936. Rapport sur la session générale de la Societe
Mycologique de France, tennue a Barcelone de 19 a 27 octubre 1935. Bull.
Soc. Mycol. Fr. 52:17-32.
Mendaza, R. and G. Diaz. 1987. Las setas. Ed. Iberduero, Vizcaya. 932 pp.
Molina, R. and J.G. Palmer. 1982. Isolation, maintenance, and pure culture
manipulation of ectomycorrhizal fungi. In: Methods and principles of
mycorrhizal research. Edited by N.C. Schenck. Amer. Phytopathol. Soc.,
St. Paul, MN. pp. 115-129.
247,
Molina, R. and J.M. Trappe. 1982. Patterns of ectomycorrhizal host specificity
and potential among Pacific Northwest conifers and fungi. For. Sci. 28:423-
458.
Moreno, G., R. Galan and A. Ortega. 1986. Hypogeous fungi from continental
Spain. |. Cryptogam. Mycol. 7:201-229.
Moreno, G., R. Galan and A. Montecchi. 1991. Hypogeous fungi from peninsular
Spain. Mycotaxon 42:201-238.
Ortega, A., R. Galan and M. de la Torre. 1981. AportaciOn al estudio de los
hongos de Andalucia. IV. El Orden Pezizales en la provincia de Granada.
Bol. Soc. Micol. Cast. 6:39-50.
Parke, J.L., R.G. Linderman and C.H. Black. 1983. The role of ectomycorrhizas
in drought tolerance of Douglas-fir seedlings. New Phytol. 95:83-95.
Singer, R. 1947. Champignons de la Catalogne. Espéces observées en 1934.
Collect. Bot. (Barc.) 1:199-246
Svrcek, M. 1958. Ill. Rad Hymenogastrales - Hlizotvaré. /n: A. Pilat, Flora CSR,
Gasteromycetes. Prace Cesk. Akad. Véd, Prague. pp. 121-208.
Tabares, M. and A. Rocabruna. 1987. Aportaci6n al conocimiento de los hongos
de la sierra de Collcerola (Catalunya). Butll. Soc. Cat. Micol. 11:83-98.
Trappe, J.M. 1977. Selection of fungi for ectomycorrhizal inoculation in nurseries.
Annu. Rev. Phytopathol. 15:203-222.
Trappe, J.M., M.A. Castellano and N. Malajczuk. 1992. Labyrinthomyces,
Dingleya and Reddellomyces gen. nov. (Ascomycotina). Aust. Syst.
Bot. 5:597-611.
Unamuno, L.M.. 1941. Enumeraci6n y distribuciOn geografica de los
Ascomycetes de la Peninsula Ibérica y de las Islas Baleares. Mem. R.
Acad. Cienc. Exactas Fis. Nat. Madr. Ser. Cienc. Nat. 8: 1-403.
Ure, D.C. and C. Maser. 1982. Mycophagy of red-backed voles in Oregon and
Washington. Can. J. Zool. 60:3307-3315.
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MY COTAXON
Volume XLVI, pp. 219-228 April-June 1993
THREE XYLARIACEOUS FUNGI WITH
SCOLECOSPOROUS CONIDIA
Y.-M. Ju
Department of Plant Pathology
Washington State University
Pullman, Washington 99164-6430
and
Felipe San Martin Gonzalez
Fac. de Agronomia
de la Universidad Aut6noma de Tamaulipas
Centro Universitario Adolfo Lopez Mateos
Cd. Victoria, Tam., 87000 Mexico
and
Jack D. Rogers
Department of Plant Pathology
Washington State University
Pullman, Washington 99164-6430
ABSTRACT
The scolecosporous anamorphs of Lopadostoma turgidum, Creosphaeria
sassafras comb. nov., and a fungus close to "Nummuzlaria" viridis are described and
illustrated from culture. Both L. turgidum and "Nummularia" Sp. produce slimy
conidia in definite conidiomata. Creosphaeria sassafras produces slimy conidia from
mycelia and from stromatic structures. The placement of these fungi in the
Xylariaceae and their resemblance to certain Diatrypaceae is discussed.
Key words: anamorph, Creosphaeria, Diatrypaceae, Hypoxylon, Lopadostoma,
Xylariaceae
One of the hallmarks of taxa of family Xylariaceae is the type of
hyphomycetous anamorph that features dry amerosporous conidia produced
holoblastically from sympodially proliferating conidiogenous cells (Rogers, 1979;
1985). Hyphomycetous genera associated with various taxa include Nodulisporium
Preuss, Geniculosporium Chesters & Greenhalgh, Lindquistia Subram. &
Chandrashekara, Xylocladium Sydow and Acanthodochium Samuels, Rogers, &
Nagasawa. We report herein on three xylariaceous fungi that, in culture, produce
more or less slimy scolecosporous conidia holoblastically from sympodially
proliferating or sympodially and percurrently proliferating conidiogenous cells.
220
Two of them produce conidia in conidiomata. These taxa do not seem to be
especially closely related to each other---indeed representing different genera---but
seem even further removed from taxa with dry conidia. We believe that these
fungi represent a distinct and heretofore unrecognized evolutionary line in the
Xylariaceae, as will be discussed later herein.
Lopadostoma turgidum
Lopadostoma turgidum (Pers.:Fr.) Traverso was discussed by the Tulasne
brothers (1863) (as Sphaeria turgida Pers.) who recognized that the valsoid
configuration of the stroma is deceptive and remarked that the ascospores
resemble hypoxyloid (xylariaceous) genera. They did not detect spermogonia, i.e.
conidiomata. Nitschke (1867) described the conidial state of L. turgidum (as
Anthostoma) as consisting of multiloculate spermagonia with cylindrical curved
hyaline spermatia produced acrogenously from densely caespitose short simple
sterigmata. Elsewhere, he gave dimensions of conidia as 10 x 1.5 “wm (1867).
Traverso (1906) cited Cytospora turgida (Nitschke) Traverso as the pycnidial state
of L. turgidum.
We received a collection of Lopadostoma turgidum from Francoise
Candoussau. This material seemed typical of the species except that the
ascospores, (7.5-)8-11.5 X 4-4.5 um, ranged shorter than reported by some authors
and more narrow than reported by most authors: 8-12 X 7-8 um (Arx & Miller,
1954); 10-13 X 5-6 zm (Dennis, 1978); 8-12 X 7-8 Wm (Ellis & Everhart, 1892);
10-13 X 5-6 “4m (Munk, 1957); 8-12 X 4-7 um (Nitschke, 1867); 8-12 X 6-8 um
(Traverso, 1906); 8-12 X 7-8 Am (Winter, 1887). Interestingly, Saccardo (1908)
described L. turgidum var. minus Sacc. (as Anthostoma) with ascospores 8-9 x 4.5-5
fim from Quercus and Castanea in France. It is possible that the present fungus
represents this taxon. We have cultured the fungus. The anamorph is illustrated
in Fig.1 C, D and described, as follows.
Colonies on 2% Difco Oatmeal Agar incubated at ca 20°C under 12 hr
fluorescent light not reaching the edge of Petri dish in 2 months, at first white,
velvety, azonate, with entire or slightly lobed margins; becoming pale luteus,
overlain with sporadic cottony mycelial masses usually associated with clear
exudates and condiogenesis. Reverse uncolored. Conidiomata convoluted or »
multiloculate with black wall ca. 0.2 mm thick, composed of prosenchymatous
elements 2.5-3 44m broad encrusted with black material. Conidiophores di- or
trichotomously branched, with yellowish stipes that become paler and then hyaline
upwards, smooth-walled, arranged in dense palisades. Conidiogenous cells
terminal, cylindrical, 15-30 X 1.5-2 zm, smooth-walled, bearing inconspicuous
conidial secession scars. Conidia produced holoblastically in sympodial sequence.
Conidia hyaline, orange in slimy masses, smooth, strongly curved to semicircular,
22-29 X 1.2-1.5 fim, (semicircular conidia 12-20 4m measured between the ends)
Fig. 1. "Nummularia" sp. and Lopadostoma turgidum. Aand B. "Nummularia’ sp.
A. Conidiophores and conidiogenous cells. B. Conidia. C and D. Lopadostoma
turgidum. C. Conidia. D. Conidiophores and conidiogenous cells.
222
with flattened bases ca. 0.5 4m broad indicating former points of attachment to
conidiogenous cells. Conidia not germinating in culture or in germination trials.
Culture deposited in American Type Culture Collection.
SPECIMENS EXAMINED: AUSTRIA. Niederdonau, Rotgraben bei Klosterneu-
burg, V.1939, Petrak F. "Mycotheca generalis," wood of Fagus silvatica, as
Lopadostoma turgida (WSP 18268); FRANCE. Carcassonne, Montreal, 8.XII.1991,
Candoussau, F. 231-A, on bark of Quercus ilex (IDR) (COLLECTION FROM
WHICH CULTURE IS DESCRIBED); Carcassonne, Montreal, 8.XII.1991,
Candoussau, F. 231, on bark of Quercus ilex (IDR) (LARGE-SPORED FUNGUS
CULTURED, BUT NOT DESCRIBED IN DETAIL). MEXICO. Tamaulipas,
Victoria municipality, "El Madrono," IX.1992, San Martin, F. 1487, on wood of
Quercus (ITCV).
Another fungus was also sent by Candoussau and was cultured. It greatly
resembled L. turgidum except that the ascospores ranged longer, (10.5-)11-16.5 X
4.5-5 um. This fungus is a variant of L. turgidum, another species buried in
Anthostoma, or a new species. In any case, cultures were similar to the above-
described except that they became blackish and incipient conidiomata did not
produce conidia.
Creosphaeria sassafras
Theissen (1910) erected Creosphaeria Theiss. based upon Creosphaeria
riograndensis Theiss. for taxa with soft light-colored stromata and brown ascospores,
remarking that the genus--"Rosellinae proximum--". Petrak (1923) believed
Anthostoma to be closely related to Creosphaeria. Later, Petrak (1951) recognized
that Creosphaeria riograndensis is conspecificwith Hypoxylon sassafras (Schwein.:Fr.)
M. A. Curtis. Miller (1961) put C. riograndensis into synonymy with H. sassafras,
the nomenclaturally much older taxon.
Martin (1967) cultured Hypoxylon sassafras, reporting the conidia to be
oval to subglobose, 1.7-3.7 X 1.7-2.7 4m. Petrini and Miller (1986), however,
reported and illustrated the anamorph from culture as belonging to Libertella
Desm., describing the conidia as crescentic, hyaline, one-celled, 13-26 X 1-2 ym.
We have recently cultured the fungus from European and Asian material. Our
results agree essentially with those of Petrini and Miller (1986) except that we
have observed conidia on stromata as well as associated with rudimentary
conidiomata in the aerial hyphae. Because this type of anamorph differs from
those of most Hypoxylon species we herein resurrect Creosphaeria, based upon
Sphaeria sassafras Schwein. It is noteworthy in this respect that Petrak (in Sydow
& Petrak, 1922) erected Creosphaeria pinea Petrak for a fungus widely accepted
as Hypoxylon diathrauston Rehm [= Rosellinia diathrausta (Rehm) L. Petrini]. The
-anamorph of this fungus is of the typical xylariaceous type (Petrini and Muller,
1986). We do not accept it as a Creosphaeria. According to Miller (1961), Rick
erected Creosphaeria verruculosa Rick. We have not seen material of this fungus
and, thus, have no opinion on its placement.
223
Creosphaenia sassafras (Schwein.: Fr.), comb. nov.
BASIONYM: Sphaeria sassafras Schwein., Schr. Nat. Ges. Leipzig 1:36. 1822.
= Sphaeria sassafras Schwein.: Fr., Syst. Mycol. II, p. 343. 1823.
= Hypoxylon sassafras (Schwein.: Fr.) M. A. Curtis, Geol. & Nat. Hist. Survey N.
C. pt. II, p. 140. 1867.
= Sphaeria callostroma Schwein., Jour. Acad. Nat. Sci. Phila. 5:15. 1825.
= Hypoxylon callostroma (Schwein.) Berk., Grevillea 4:51. 1875.
= Hypoxylon bifrons De Not., Sphaer. Ital. Cent. I, p. 18. 1863.
? = Sphaena prinicola Berk. & M. A. Curtis, Grevillea 4:142. 1876.
= Rosellinia prinicola (Berk. & M. A. Curtis) Sacc., Syll. Fung. I, p. 263.
1882.
? = Rosellinia linderae Peck, 49th Rep. N. Y. State Mus., p. 24. 1896.
= Hypoxylon valsarioides Speg., Fungi Chilenses, p. 48. 1910.
= Creosphaeria niograndensis Theiss., Beih. Bot. Centralb. 27:396. 1910.
? = Anthostomella sinensis Teng & Ou, Sinensia 7:198. 1936.
The teleomorph is as described by Miller (1961) and Petrini & Miiller
(1986). The anamorph is illustrated in Fig. 2 A-E and described as follows.
Colonies on 2% Difco Oatmeal Agar incubated at 20°C under 12 hr
fluorescent light covering Petri dish in 4 wk, at first white, felty, azonate, with
diffuse margins and pinkish diffusible pigment, becoming overlain with a blackish
gray layer of floccose hyphae. Reverse uncolored. Stromata formed in old
_ cultures, scattered, appressed cushion-shaped, 2-5 mm diam x ca. 1 mm high, black.
Conidium-bearing regions on surfaces of both mycelium and stromata, or in
mycelial cavities. Conidiophores dichotomously branched, with brown stipes that
become paler upwards, smooth-walled, loosely arranged when arising from aerial
hyphae but in dense palisades when arising from stromata and mycelial cavities.
Conidiogenous cells terminal, cylindrical, 10-22 x 2-3 4m, smooth-walled, often
bearing denticulate conidial secession scars, infrequently with several annellations.
Conidia produced holoblastically in sympodial sequence or from percurrently
proliferating conidiogenouscells. Conidia hyaline, wet to somewhat slimy, smooth,
strongly curved, (13-)15-22 x 1.5-2 44m, with flattened bases ca. 0.5 4m broad
indicating former points of attachment to conidiogenouscells. Conidia germinating
in culture.
Culture deposited in American Type Culture Collection.
SPECIMENS EXAMINED: BRAZIL. Sao Leopoldo, 1909, leg. Theissen, in
Rick, J. "Fungi Austro-Americani" no. 316, on corticated wood, as Creosphaeria
nograndensis Theiss. (BPI; TYPE). CHILE. no. 1965. Microscope slide of Shear,
C. L. Labelled as type of Hypoxylon valsaroides Speg. (BPI). FRANCE.
Bugangue, Oloron, 2.11.1992, Candoussau, F., Fagus (IDR) (CULTURED).
ITALY. Microscope slide of Shear, C. L. bearing a portion of stroma and
ascospores. Labelled as type of Hypoxylon bifrons De Not. from herb FI (BPI);
TAIWAN. Nan-Tou County, Huei-Sun, 22.1V.1985, Ju, Y.-M., NTU-74042201,
fallen corticated twigs (JDR; NTU); Taipei City, Young-Ming-Shan, 8.1.1988,
Chang, T.-M., NTU-77010801, corticated wood (JDR; NTU) (CULTURED).
224
UNITED STATES. Pennsylvania, Bethlehem, Collins Coll. no. 39, Syn. 1208, as
Sphaeria sassafras LVS. (PH; TYPE); Pennsylvania, Bethlehem, Collins Coll. no. 33,
Syn. 1200, as Sphaeria sassafras LVS. (PH;TYPE). Pennsylvania, Bethlehem,
Collins Coll. no. 39, Syn. 1208, as Sphaeria callostoma LVS (PH;?7 HOLOTYPE).
The anamorph of Creosphaeria sassafras, producing elongated conidia from
conidiogenous cells that proliferate sympodially or percurrently, resembles that of
Hypoxylon microplacum (Berk. & M. A. Curtis) J. H. Miller (Glawe & Rogers,
1986). Conidia of the latter species tend to be dry.
"Nummularia" sp.
Theissen (1908) described Nummularia viridis Theiss. on material collected
in Brazil. Stromata were described as "--flavo-viridula, dein viridi-fuliginea margine
laetiore flavidulo--.". Miller (1961) put N. viridis into synonymy with Hypoxylon
hypophlaeum (Berk. & Ravenel) J. H. Miller. We were unable to locate type or
other material of N. viridis in the USA or herb. K. We examined a specimen of
Rick from herb. S that is from the type locality, but is probably not type material.
It is distinct from H. hypophiaeum. In any case, we have cultured a fungus that
seems close to N. viridis as described by Theissen (1908). It is briefly described,
as follows: Stroma applanate, 5 cm x 2 cm x <1 mm thick, smooth, cracked in the
manner of Diatrype stigma. Exterior greenish tan, blackish beneath, but white
around perithecial bases which are embedded directly in the substrate. Texture
waxy. Perithecia 0.1-0.2 mm diam. Ostioles umbilicate, slightly raised, blackish.
Asci short- to long-stipitate, with ascospores arranged in uniseriate or biseriate
manner, ca. 90 x 5-10 wm, the spore-bearing part ca. 40 wm long, with minute
plate-like ascus apical ring bluing in Melzer’s iodine reagent. Ascospores brown,
smooth, one-celled, ellipsoid-inequilateral, 8-10.5 x 3.5-4.5 um, with long straight
germ slit on concave part. It differs from the Rick fungus which has more discrete
uncracked stromata that lack green tones, has raised black ostioles, and has
ascospores that are more narrow, i.e. 3-3.5 wm.
Our fungus might or might not be conspecific with N. viridis. It certainly
represents a new genus, based upon its elongated wet conidia borne in conidiomata
and teleomorphic features, but we are not describing it as such until a broader
search is made for material of N. viridis which might form the type of a new genus.
The anamorph is illustrated in Fig. 1 A, B and described, as follows.
Colonies on 2% Difco Oatmeal Agar incubated at 20°C and 12 h of
fluorescent light covering Petri dish in 5 wk, at first yellow, velvety, with diffuse
margins, becoming greenish olivaceous, zonate. Reverse brownish. Conidium-
bearing regions distributed on concentric zones, within or upon a stroma. Stroma
superficially greenish, black beneath, slightly carbonaceous, composed of spheroid,
dark, thick-walled cells 3-4.5 44m diam, overlying white prosenchymatous layer.
Fig. 2. Creosphaeria sassafras. A-D. Conidiophores and conidiogenous cells
showing conidial production and conidiogenous cells proliferating sympodially
and/or percurrently. E. Conidia.
226
Conidiophores di- or trichotomously branched, with hyaline to pale brown stipes,
smooth-walled, arranged in dense palisades. Conidiogenous cells terminal,
cylindrical, smooth, 25-40 x 1.5-2 am, bearing slightly denticulate secession scars.
Conidia produced holoblastically in sympodial sequence. Conidia hyaline, orange
in slimy masses, smooth, slightly curved, 10-17 x 1.2-1.8 zm, with flattened bases
ca. 0.5 um broad indicating former points of attachment to conidiogenous cells.
Conidia not germinating.
Culture deposited in American Type Culture Collection.
SPECIMENS EXAMINED: BRAZIL. Nova Petropolis, Rick, J. 226, as
Nummularia grisea Rick (BPI; TYPE); Sao Leopoldo, 1906, Rick, J., as Hypoxylon
grisea Rick (BPI). Sao Leopoldo, III.1908, Rick, as Nummularia viridis Theiss. (S).
MEXICO. Nuevo Le6n, Santiago, El Cercado, 16.1V.1988. San Martin, F. #428,
on Quercus, as "Nummularia" sp. (ITCV;JDR) (CULTURED). USA. Louisiana,
St. Martinsville, 28.III.1890, Langlois, "Fungi Ludoviciana: no. 2278, as Hypoxylon
cinereum Ellis & Everh. (NY,TYPE); New Jersey, Newfield, 1V.1894, Ellis, J. B.
"Fungi Columbiana" 429, on Magnolia glauca, as Nummulana hypophloea (Berk.
& Ravenel) Cooke (WSP 4783); South Carolina, Monck’s Corner, 26.X.1923, as
Hypoxylon hypophlaeum (Berk, & Ravenel) J. H. Miller (BPI); South Carolina,
1855, Ravenel, "Fung. Carol., Fasc. 4, no. 38, as Diatrype hypophloea Berk. &
Ravenel (BPI).
The teleomorphic stroma of this fungi resembles that of Hypoxylon
microplacum. The conidia likewise resemble those of H. microplacum, but those
of this latter fungus are dry and not formed in conidiomata. A search of
herbarium specimens did not settle the identity of the fungus that we cultured. It
did, however, reveal that Hypoxylon grisea Rick, based on the type, and another
Brazilian specimen cited as H. grisea are neither conspecific nor to be equated with
H. hypophlaeum as Miller (1961) believed. Likewise, examination of type material
of Hypoxylon cinereum revealed that it is probably a taxon distinct from H.
hypophlaeum. One might be tempted to remove all of these taxa to Biscogniauxia
O. Kuntze on the basis of teleomorphic features. It is suspected, however, that
cultures could reveal anamorphic differences that would influence generic
assignment. We thus continue to use old and untenable generic names.
DISCUSSION
The fungi described herein appear to represent an evolutionary line in the
Xylariaceae, based primarily on the anamorphs. Scolecosporous conidia are
unusual among cultured Xylariaceae. Wet to slimy conidia borne in or on
conidiomata represent a striking contrast to the dry-spored hyphomycetous states
of most investigated xylariaceous fungi. Interestingly, the fungi described here--
although possessing cardinal xylariaceous features such as brown ascospores with
germ slits and amyloid ascus rings--differ developmentally among themselves.
Lopadostoma turgidum develops in a more or less valsoid manner within bark and
wood. Creosphaeria sassafras appears to develop in the manner of the colored
Hypoxylon species, i.e. as an erumpent stroma that develops perithecia in the
227
periphery following conidial production on the stroma. "Nummularia" sp.
apparently has a bipartite stroma of the type associated with Biscogniauxia or
Camillea Fr. (Laess¢e et al, 1989). The relationships among these three taxa are
presently unclear.
The anamorph of Creosphaeria sassafras has a resemblance to that of
Hypoxylon microplacum in conidial shape, in having both sympodial and percurrent
conidiogenous cell proliferation, and in that conidia of both species germinate to
some extent. The teleomorphs, however, are developmentally quite different, that
of H. microplacum being of the Biscogniauxia type. It is currently uncertain if H.
microplacum should be removed to Biscogniauxia or placed in a new genus.
The anamorphs of the three species described here are reminiscent of the
Diatrypaceae in the more or less slimy scolecosporous conidia produced on
conidiogenous cells that proliferate sympodially and/or percurrently and in the
production of more or less distinct conidiomata (see discussion by Glawe &
Rogers, 1986). The conidia of Lopadostoma turgidum and "Nummularia" sp. have
not been seen to germinate. Conidial germination has not been observed in most
diatrypaceous fungi and, when observed, occurs in very low numbers (Ju et al,
1991). The conidial nucleus of many xylariaceous and diatrypaceous fungi occupies
much of the volume of the spore (Glawe and Rogers, 1986; Roe, 1974). This is
true for conidia of "Nummularia" sp., the only species examined cytologically.
We do not yet know the relationships of the Xylariaceae with the
Diatrypaceae. In 1986 Glawe and Rogers wrote that the Diatrypaceae and the
Xylariaceae probably arose from a common ancestor. It was stated therein that
the Xylariaceae depend exclusively on air-dispersed conidia (Glawe & Rogers,
1986). Obviously, our impression of the Xylariaceae is now amended to include
taxa with water-dispersed conidia that, in some cases, are borne in (or on)
conidiomata. The interfaces of the Xylariaceae and Diatrypaceae remain unclear.
ACKNOWLEDGEMENTS
PPNS. No. 0136, Department of Plant Pathology, Project 1767, Washington
State University, Agricultural Research Center. This study was supported in part
by National Science Foundation grant BSR-9017920 to JDR and a CONACYT
grant to FSMG. We thank the curators of herbaria BPI and NY for specimens.
We are grateful to Francoise Candoussau, Pau, France for fresh collections. We
thank Lori Carris and Jane Lawford, Washington State University, respectively, for
criticizing the manuscript and typing the manuscript. We thank J. L. Crane,
University of Illinois, for reviewing the manuscript.
LITERATURE CITED
Arx, J. A. von & E. Miller. 1954. Die Gattungen der amerosporen
Pyrenomyceten. Beitr. Kryptogamenfl. Schweiz 11:1-434.
Dennis, R. W. G. 1978. British Ascomycetes. J. Cramer. Vaduz. 385 p.
Ellis, J. B. & B. M. Everhart. 1892. North American pyrenomycetes. Newfield,
New Jersey. 793 p.
Glawe, D. A. & J. D. Rogers. 1986. Conidial states of some species of
228
Diatrypaceae and Xylariaceae. Can. J. Bot. 64:1493-1498.
Ju, Y.-M., D. A. Glawe, and J. D. Rogers. 1991. Conidial germination in Eutypa
armeniaceae and selected other species of Diatrypaceae: Implications for
the systematics and biology of diatrypaceous fungi. Mycotaxon 41:311-320.
Laessge, T., J. D. Rogers, and A. J. S. Whalley. 1989. Camillea, Jongiella and
light-spored species of Hypoxylon. Mycol. Res. 93:121-155.
Martin, P. 1967. Studies in the Xylariaceae IV. South African and foreign
species of Hypoxylon sect. Entoleuca. J. S. African Bot. 34:153-199.
Miller, J. H. 1961. A monograph of the world species of Hypoxylon. Univ. of
Georgia Press, Athens. 158 p.
Munk, A. 1957. Danish pyrenomycetes. Dansk Botan. Arkiv. 17:1-491.
Nitschke, T. 1867. Pyrenomycetes Germanici. Breslau. 320 p.
Petrak, F. 1923. Mykologische Notizen. 6. Ann. Mycologici 21:182-335.
Petrak, F. 1951. Engebnisse einer Revision der Grundtypen vershiedener
Gattungen der Askomyzeten und Fungi Imperfecti. Sydowia 5:328-356.
Petrini, L. E. & E. Miller. 1986. Teleomorphs and anamorphs of European
species of Hypoxylon (Xylariaceae, Sphaeriales) and allied genera.
Mycologia Helvetica 1:501-627.
Roe, G. W. 1974. Fine structural studies on the asexual state of Xylaria and
related genera. Ph.D. dissertation (2 volumes). Univ. of Liverpool.
Rogers, J. D. 1979. The Xylariaceae: Systematic, biological, and evolutionary
aspects. Mycologia 71:1-42.
Rogers, J. D. 1985. Anamorphs of Xylaria: Taxonomic considerations. Sydowia
38:255-262.
Saccardo, P. A. 1908. Notae Mycologicae. Annales Mycologici 6:553-569 + Tab.
XXIV.
Sydow, H. & F. Petrak. 1922. Ein Beitrag zur Kenntnis der Pilzflora
Nordamerikas, insbesondere der nordwestlichen Staaten. Annales
Mycologici 10:178-218.
Theissen, F. 1908. Novitates riograndes. Ann. Mycologici 6:341-352.
Theissen, F. 1910. Fungi riograndenses. Botanisch. Centralblatt Beihefte 27:384-
411.
Traverso, J. B. 1906. Flora Italica Cryptogama. I: Fungi. Pyrenomycetae.
Societa Botanica lialiana. R. S. Casciano. 325 pp.
Tulasne, L. R. & C. Tulasne. 1863. Selecta fungorum carpologia. Vol. II. Paris.
(English translation: W. B. Grove, 1931. Oxford University Press,
Oxford).
Winter, G. 1887. Ascomyceten: Gymnoasceen und Pyrenomyceten. IN: L.
Rabenhorst, ed. Kryptogamen-Flora von Deutschland, Oesterreich und
der Schweiz. Abtheilung 2. 2nd ed. E. Kummer, Leipzig. 925 p.
MYCOTAXON
Volume XLVII, pp. 229-258 April-June 1993
BISCOGNIAUXIA AND CAMILLEA IN MEXICO
Felipe San Martin Gonzalez!
Instituto Tecnolégico de Ciudad Victoria
Ap. Post. no. 175
Cd. Victoria, Tamaulipas, Mexico
and
Jack D. Rogers
Department of Plant Pathology
Washington State University
Pullman, WA 99164-6430
ABSTRACT
Thirteen taxa of Biscogniauxia and fourteen taxa of Camillea from Mexico
are considered and included in keys. Cultural and anamorphic data are presented
for some taxa. A new combination, Biscogniauxia fuscella, is made. The new
species--Camillea guzmanii, C. magnifica, and C. mexicana--are erected.
Key words: Biscogniauxia, Camillea, Nodulisporium, Xylariaceae, Xylocladium
In 1989 San Martin Gonz4lez and Rogers published a preliminary account
of Xylaria Hill ex Schrank in Mexico based primarily on collections made by the
former (hereinafter called San Martin) prior to 1988. In 1988 San Martin spent
several months of extensive collecting in 34 principal locations in 12 Mexican states.
Most of the collection locations were in the eastern, southeastern, and southern
States. A detailed list of collection locations is appended to this paper. This paper,
and those to follow that will deal with other genera, include data from all of San
Martin’s collecting activities.
Mexico is a geographically, geologically, climatically, and_floristically
complicated country. This is reflected in its mycoflora. References such as
Rzedowski (1978) and Koeppen (1948) are indispensable to the understanding of
‘Present address: Fac. de Agronomia de la Universidad Aut6noma de
Tamaulipas, Centro Universitario Adolfo Lopez Mateos, Cd. Victoria, Tamaulipas
87000, Mexico
230
the physical aspects and vegetation, and the climate, respectively, of Mexico. As
one might suspect, mycofloristic elements of the United States, Central America,
and South America are found there. Where appropriate, comments on distribution
are included in the "NOTES" for each taxon.
MATERIALS AND METHODS
Collected materials were air-dried and brought in culturable condition to
Washington State University under appropriate permits issued by APHIS
authorities in Hyattsville, Maryland. Teleomorphic material was placed in ITCV
and JDR, the personal herbarium of J. D. Rogers. Herbarium designations follow
Holmgren, et al. (1990).
Cultures were made from teleomorphic stromata in the following way. The
upper part of a perithecium-bearing stroma was removed with a sterile razor blade.
The content of the exposed perithecium was rehydrated with sterile distilled H,O,
scooped out with a sterile needle and transferred to a Petri plate containing 2%
Difco potato dextrose agar amended with 5g/L of Difco yeast extract (PDYA).
Following germination of ascospores, hyphal tips were severed and transferred to
2% Difco Oatmeal agar (OA) in 9 cm diam Petri plates and incubated under
laboratory conditions of approximately 12 hr per day fluorescent light at 20°C.
Cultures thus were considered as originating from multiple ascospores. Dried
cultures were deposited in ITCV and WSP.
Conidial structures were examined by a technique involving coverslips coated
with mucilage (Callan & Rogers, 1990). Anamorphic and teleomorphic structures
were initially examined and measured in water. Structures were examined by
brightfield (BF) microscopy, differential interference contrast (DIC) microscopy,
and scanning electron microscopy (SEM). SEM techniques were as used by
Rogers (1977).
Biscogniauxia O. Kuntze
Biscogniauxia largely includes lignicolous taxa that were traditionally included
in the illegitimate genus Nummulanra Tulasne or, more recently, in Hypoxylon Bull.
(Miller, 1961; Pouzar, 1979), Stromata are applanate, more or less cylindrical, or
cupulate. Stromata are usually (probably always) bipartite. An outermost layer
that at first covers and occludes the ostioles is shed prior to perithecial maturity,
exposing the ostioles. In cases where stromata are formed within bark, the
outermost layer adheres to overlying bark when it is ruptured by the growing
stroma and thrown off. Ascospores are usually brown to nearly black, smooth-
walled, often nearly equilateral, possess a germ slit, and lack a dehiscent perispore.
They are one-celled or, in some species, two-celled with one of the cells hyaline.
The apical ascus ring is usually amyloid (bluing in Melzer’s iodine reagent) and
usually is broader than high. In some species the ring is reduced to a thin flat
perforated plate. Known anamorphs of Biscogniauxia are accommodated in
Nodulisporium Preuss.
231
There are among the included taxa some species or varieties that have not
formally been transferred to Biscogniauxia. In some cases, there is substantial
doubt about the actual taxonomic disposition. This is particularly true with the
complex of taxa surrounding Biscogniauxia nummularia, some of which were
considered varieties of this species (as Hypoxylon nummularium) by J. H. Miller
(1961).
iB
KEY TO SPECIES OF BISCOGNIAUXIA FROM MEXICO
Stromata with whitish-gray to white surface at maturity ............ 2
Stromata with dark brown, blackish to shiny black surface at maturity . 4
2. Perithecial ostioles umbilicate to slightly raised. Ascospores
brown to dark brown with one or both ends narrowly or broadly
rounded, one end often bearing a minute appendage, 13.5-16
(-16.5) (including appendage) x 7-8 4m. On Celtis laevigata......
sles ae ats uate: oth LanCel oetoych ofletal »/% telat ho sL bbe Biscogniauxia fuscella
Zz, Perithecial ostioles papillate. Ascospores larger............. 3
Ascospores (30.5-)32-34(-37) x (12-)13-15 wm. On Quercus spp. .......
PET CRA ne er Ue eBO nti c Turare pst Biscogniauxia atropunctata
Ascospores 18.5-20(-21) x (7-)7.5-9 am. On Quercus sp. .........+.5-
hs eaten Se ey eri id tm Baan tb Biscogniauxia sp. (José Castillo 1100B)
4. Ascospores brown to dark brown with one end rounded and the
other truncate or bearing a hyaline to pale brown cellular
AD PCNA GE ee tteecare ie Chet take fo) -Nehe sue ese e is tort tay Gey sissies aie, 2 5)
4. Ascospores devoid of cellular appendage or notably truncate
SUITUE Aad Fee eee oe Te Oe Coc CEL ARE GD USM ete WAR sh oe Chan ve Rae 6
Perithecial ostioles indistinct. Ascospores (19-)21-24(-25) (not including
appendage) x (10-)11-14.5(-15) zm, with germ slit straight running the entire
length of the dark spore body ... Biscogniauxia divergens var. macrospora
Perithecial ostioles umbilicate in more or less deep pits. Ascospores (10-)11-
13 (not including appendage) x 6-7.5(-8) 4m with germ slit straight running
3/4 ofthe: darkispore body v.t-).:) iu) sas Ge Biscogniauxia grenadensis
Gite WASCOSDOTES LIGHT DTOWM tas) tale Biolog lajitys ay beet eee ain, + Tele tle fate i,
6. Ascospores brown, dark brown to almost black ............. 8
Perithecia 1-1.5 mm high x 1-1.7 mm broad. Ascospores ellipsoid with
rounded to narrowly-rounded ends, (13-)14-19(-21) x 5-7 4m .........
or ee rg ra lc ote, oe En NE ar gebrarS fge Len Hypoxylon cf. comedens
232
ds Perithecia 0.1-0.2 mm broad. Ascospores ellipsoid-inequilateral to navicular
with narrow ends, 5-6 x (2-)2.5(-3) um ....... Hypoxylon microplacum
8. Stromatal surface shiny black. Perithecial ostioles prominent,
conic-papillate. Ascospores dark brown to almost black, 15-
T7DS(-19X 1-SEO) Ny ee oe aha er Biscogniauxia mediterranea
8. Stromatal surface dark brown to dull black. Ostioles not
prominently conic-papillatety in. ce aie ree ote erties wicks 9
p: Ascospores ellipsoid-inequilateralto navicular with narrow to acute ends, 13-
16(-17) x (4.5-)5-5.5(-6) zm, with straight germ slit much less than spore-
ISTO, eta hte peter ee let eee Biscogniauxia sp. (San Martin 725)
2; Ascospores with rounded or narrow ends, with germ slit running the full
spore-length or nearly:sO > ya has 2 ss a vente, ee ty eee 10
10. Ascospores subspherical with rounded ends, dark brown to
almost black, (10-) 11-12 (-13) 227-89) ema ee
iets clef, 6 keh LAR gees octane. ele, eit Biscogniauxia cf. nummularia
NOTE: Ascospores of Mexican material of B. nummularia frequently have
pinched ends.
10. Ascospores ellipsoid-inequilateral to navicular ............. 11
11. Ascospores with narrow to narrowly-rounded ends, dark brown to almost
black, (13-)13.5-17(-19) x 8-8.5 um. Stromata on Quercus spp. wood in
temperate forests}. ar. ee Biscogniauxia nummularia var. exutans
11. Ascospores smaller. Stromata on wood in subtropical forests or in tropical
FALUNMOTESIS (eee ere ean ne IV Wicenp dare occ eaters Pee car ek ae eee ne 12
12,» 5 -AsCospores: (8.54)9-12)x9.5:6(-6.5)) [Mens = eae ee te ne
SCO CAEP CTR EME ey RE Biscogniauxia nummularia var. merrillii
12s), Ascospores.10212.5(-13.5)sx.6.5-7(-8), Me se. ee ee
I A ee Ste Biscogniauxia nummularia var. "pseudopachyloma"
Biscogniauxia atropunctata (Schwein.) Pouzar, Cesk4 Mykol. 33:216. 1979.
The teleomorph is much as described by Miller (1961). Ascus apical ring
bluing in Melzer’s iodine reagent, cuneate, 3-3.5 wm high x 7-8 “wm broad.
Ascospores (30.5-)32-34(-37) x (12-)13-15 ym, with straight germ slit full-length on
convex side. The anamorph was described by Martin (1969) (as Numulariola
atropunctata).
SPECIMEN EXAMINED: Nuevo Leén state, Iturbide municipality, Ejido La
Purisima, Alt. 2000 m, no date, Marmolejo 185, on wood of Quercus sp. in an oak
forest (ITCV;JDR).
233
NOTES: This pathogen of Quercus spp. principally in the southern USA (Miller,
1961), was found in México on its expected host genus.
Hypoxylon cf. comedens Ces., Atti Accad. Sci. Fis. 8:19. 1879.
Teleomorph is as described by Miller (1961) except that the thickness of the
stroma is greater, i.e., 1.8-4 mm vs 1-1.5 mm, and perithecia are broader, i.e., 1-2.0
mm vs 1-1.2 mm.
SPECIMEN EXAMINED: Chiapas state, Ocosingo municipality, Ejido Boca de
Chajul, VII.87, San Martin 376, on wood in evergreen tropical rain forest
(ITCV;JDR).
NOTES: This fungus is similar to Biscogniauxia in the absence of a basal
entostroma which at maturity is partially lost (as in Ustulina Tul. & C. Tul.), the
applanate and erumpent stroma, the ostiolar pits, and the light brown, smooth
ascospores. It seems most prudent, however, to await the availability of fresh
material so that further observations and cultural data can be brought to bear on
the taxonomic status of this fungus. Martin (1969) described material from Mexico
(as Numulariola). He reported cultures bearing conidiophores of the Nodulispor-
tum type (as Acrostaphylus) and of the Xylocladium type (as Basidiobotrys). This
indicates that his fungus might, in fact, be a species of Camillea. We have not seen
Martin’s material, however. Type material of H. comedens is from Indonesia and
its smooth ascospores that possibly bear a germ slit are unlike Camillea (Laessge
et al., 1989).
Biscogniauxia divergens (Theiss.) A. J. S. Whalley & T. Laessge var. macrospora (J.
H. Miller) A. J. S. Whalley & T. Laessge, Myc. Res. 94:239. 1990.
The teleomorph is much as described by Miller (1961). Ascospore germ slit
Straight, running the entire colored part on convex side.
SPECIMEN EXAMINED: Chiapas state, Ocosingo municipality, Montes Azules
Biosphere Reserve, 21.VIII.87, San Martin 400 & 400B, on dicot wood in
evergreen tropical rain forest (ITCV;JDR).
NOTES: This material is overmature, but stromatal characters, and shape and size
of ascospores fit well Miller’s (1961) description of H. divergens var. macrospora.
The type material of this variety was originally collected somewhere in
México in 1895 by C. L. Smith. This material was identified by Berkeley as
Hypoxylon tinctor (unpublished data) and later recognized as a new variety of
Hypoxylon divergens by Miller (1961).
Biscogniauxia fuscella (Rehm) San Martin & J. D. Rogers, comb. nov.
Basionym: Nummularia fuscella Rehm, Ann. Mycologici 2:176. 1904.
Hypoxylon fuscellum (Rehm) J. H. Miller, Mycologia 25:325. 1933.
Stromata applanate, erumpent, 0.7-25 cm long x 0.7-2.5 cm broad x 0.5-0.7
234
mm high; externally whitish-gray, internally black. Texture hard but fragile.
Surface smooth to undulate due to irregularities of the substrate. Perithecia ovoid,
0.4-0.7 mm broad. Ostioles impressed to slightly papillate. Asci cylindrical, eight-
spored, with uniseriate spore arrangement, very short-stipitate, 100-118 2m total
length x 9.5-12 wm broad, the spore-bearing part 92-106 zm, with apical ring
bluing in Melzer’s iodine reagent, slightly cuneate, 1.5-2 zm high x 4-4.5 wm broad.
Ascospores brown to dark brown, ellipsoid-inequilateral to navicular, with narrowly
rounded ends, or some with one end narrowly rounded and the other truncate,
13.5-16(-16.5) x 7-8 um, with straight spore-length germ slit whose margins
protrude in the manner of clam shell sutures, often with a cellular (?), hyaline
appendage at one end ca. 1.5 wm long.
Colonies covering Petri plate in 3 wk, sectored, at first white, velvety,
somewhat zonate, then with white, woolly growth overlaying the surface, with well-
defined margins. Odor strongly yeasty. Reverse at first yellow, then green.
Conidium-bearing regions over the entire surface of the colony. Conidiophores
erect, branched, with well-defined main axis, 3-4.5 4m broad, brownish toward
bases, warted. Conidiogenous cells terminal and lateral, di- or triverticillately
branched, cylindrical, 9-15 x 3-4 wm, hyaline, bearing poroid conidial secession
scars, with apices somewhat swollen due to conidial production. Conidia cream in
mass, individually hyaline, subglobose to ellipsoid, (4.5-)5-6 x 3-4 4m, with flattened
bases indicating former points of attachment to conidiogenous cell. Conidia
germinating in water.
SPECIMEN EXAMINED: Tamaulipas state, Victoria municipality, Km 2 Victoria
City-Matamoros City road, 5.1X.88, San Martin 1011 (CULTURED), on wood of
Celtis laevigata Willd. ("palo blanco") in semidesertic scrub with Pithecellobium
flexicaule (Benth.) Coult. (TCV;JDR).
ADDITIONAL SPECIMENS EXAMINED: Nummularia fuscella, USA, Texas,
Long 481, Cat. # 11202, (BPI;HOLOTYPE); Hypoxylon atropurpureum Fr. =
Hypoxylon cinereo-lilacinum J. H. Miller, USA, Iowa, Decorah, Sept. 1882, Cat.
# 1180 (BPI;ISOTYPE).
NOTES: The Mexican fungus is similar to the type of Nummulana fuscella Rehm.
The applanate stroma, the cuneate apical ring, the color of the ascospores, and the -
conidial state assignable to Nodulisponum Preuss clearly link N. fuscella to species
of Biscogniauxia. Miller (1961) had placed N. fuscella in his subsection Papillata
of section Papillata Hypoxylon due to the more or less papillate ostioles. The color
of the ectostroma, the shape of the ostioles, and the size of the ascospores of the
Figs. 1-8. Biscogniauxia sp. (José Castillo 1100B), Camillea magnifica, and
Camillea mexicana. 1,2. Biscogniauxia sp. (J. Castillo 1100B). 1. Stromata, x 3.8.
2. Stromatal surface, x 16. 3-5. C. magnifica. 3. Stroma, x 6. 4. Stromatal
surface, x 16. 5. Stroma (arrow) accompanied by other stromata of a
Biscogniauxia, x 1. 6-8. C. mexicana. 6. Portion of stroma, x 1. 7. Stromatal
surface, x 16. 8. Stromatal surface, x 24.
235
236
Mexican collection are shared by Biscogniauxia cinereo-lilacina (J. H. Miller)
Pouzar (= H. cinereo-lilacinum J. H. Miller), but the latter lacks the strong amyloid
reaction of the ascal apical ring, lacks appendaged ascospores, and apparently is
restricted to Tilia spp. in cooler localities (Miller, 1961).
Biscogniauxia grenadensis (J. H. Miller) A. J. S. Whalley & Laessge, Myc. Res.
94:239. 1990.
Teleomorph is much as described by Miller (1961). Ascospores with straight,
short germ slit in dark body of spore.
SPECIMENS EXAMINED: Chiapas state, La Trinitaria municipality, Montebello
Lagoons, 25.V.88, San Martin 840 & 848, on dicot wood in an oak forest mixed
with tropical species (ITCV;JDR); San Cristobal de las Casas municipality,
El Huitepec Hill, 4.V1.88, San Martin 727, on dicot wood in oak forest
(TCV;JDR).
NOTES: Collection San Martin 727 is young, but some mature ascospores were
found and these were typical of B. grenadensis.
Biscogniauxia mediterranea (De Not.) Kuntze, Rev. Gen. Plant. 2, p. 398. 1891.
The teleomorph is as described by Miller (1961). The anamorph was
described by Martin (1969) (as Numulariola mediterranea), and Jong & Rogers
(1972).
SPECIMEN EXAMINED: Tamaulipas state, GOmez Farias town, 14.1X.88,
San Martin 1130, on wood in median subdeciduous tropical rain forest
(ITCV;JDR).
ADDITIONAL SPECIMEN EXAMINED: Biscogniauxia mediterranea, Portugal,
1969, N-275 (JDR).
NOTES: Mexican material is typical of the species. It was collected in a tropical
rain forest.
Figs. 9-20. Biscogniauxia sp. (José Castillo 1100B), Biscogniauxia sp. (San
Martin 725), Camillea fossulata. 9-11. Biscogniauxia sp. (J. Castillo 1100B). 9.
Ascospore, x 1500. 10. Ascospore with germ slit, x 1300. 11. Ascus apical ring,
x 1400. Figs. 12-14. Biscogniauxia sp. (SM 725). 12. Ascus apical ring, x 2300.
13. Ascospore, x 2400. 14. Ascospore with short, straight germ slit on concave
side, x 2300. 15-20. C. fossulata. 15. Ascospore, x 2000. 16. Portions of two asci
with ascospores, x 1900. 17. Ascus apical ring, x 2250. 18. Ascospore with
ornamentation, x 20000. 19. Ascospore with ornamentation, x 2500. 20.
Ascospore showing ornamentation, x 11000.
Figs. 9-14 by BF. Figs. 15-17 by DIC. Figs. 18-20 by SEM. Figs. 9, 10, 13-
16 from water mounts. Figs. 11, 12, 17 from mounts in Melzer’s reagent.
237
238
Hypoxylon microplacum (Berk. & M. A. Curtis) J. H. Miller, Monogr. of the world
species of Hypoxylon, p. 129. 1961.
The teleomorph is as described by Miller (1961). The anamorph is as
described by Jong & Rogers (1972).
SPECIMENS EXAMINED: Tamaulipas state, Casas municipality, El Lajeadero
Ranch, 8.X1.86, Jesis Garcia 5824, on wood in submontane scrub (ITCV;JDR);
Gé6mez Farias town, II.21.88, Chac6n-Jiménez 240B, on wood in median
subdeciduous tropical rain forest ITCV;JDR).
NOTES: This species was reported by Pérez-Silva (1983) from the state of
Oaxaca, México. Martin (1969) considered it to be a Numulariola, but did not
observe the scolecosporous conidia reported by Jong & Rogers (1972) and Glawe
& Rogers (1986). We are tempted to move it to Biscogniauxia, but she a that
it will ultimately be placed in another genus.
Biscogniauxia nummulana (Bull.:Fr.) Kuntze var. exutans (Cooke) Van der Gucht,
Mycotaxon 45:266. 1992.
Hypoxylon nummularium Bull.:Fr. var. excutans (Cooke) J. H. Miller, Monogr. of
the world species of Hypoxyion, p. 124. 1961.
Teleomorph is much as described by Miller (1961) and Ju (1990). Asci
cylindrical, eight-spored, short-stipitate, 119-150 42m total length x 10-11(-11.5) “zm
broad, the spore-bearing part 100-120 wm, with apical ring bluing in Melzer’s
iodine reagent, cuneate, 3(-3.5) 4m high x 4 “4m broad. Ascospores dark brown to
almost black, ellipsoid-inequilateral to navicular with narrow to narrowly rounded
ends, (13-)13.5-17(-19) x 8-8.5 4am, with germ slit straight spore-length on convex
side.
The anamorph was described by Martin (1969) (as Numulariola exutans).
SPECIMENS EXAMINED: Nuevo Leon state, Iturbide municipality, Ejido La
Purisima, Alt. 2000 m, no date, Marmolejo 199, on wood of Quercus affinis Mart.
& Gal. in an oak forest (ITCV;JDR). Tamaulipas state, Hidalgo municipality,
Fjido Conrado Castillo, VII.87, San Martin 453, on Quercus sp. wood in oak-pine
forest ITCV;JDR).
NOTES: This is what Miller (1961) called Hypoxylon nummularium var. exutans.
We reluctantly accept it as a variety of Biscogniauxia nummularia because we are
uncertain if it is to be considered as a variety or as B. exutans as Martin (1969)
believed (as Numulaniola). We require cultural and other data to reach a firm
decision.
Mexican material differs from Taiwan material (Ju, 1990) in having smaller
apical rings, ie., 3(-3.5) zm high x 4 wm broad vs 5-6 4m high x 5 zm broad. The
taxonomic significance of apical ring dimensions in Biscogniauxia is unknown.
Biscogniauxia nummulana (Bull.:Fr.) Kuntze var. merrillii (Bres.) Van der Gucht,
Mycotaxon 45:276. 1992.
Hypoxylon nummularium Bull.:Fr. var. merrillii (Bres.) J. H. Miller, Monogr. of
239
the world species of Hypoxylon, p. 126. 1961.
The teleomorph is as described by Miller (1961). Colony morphology and
anamorphic structures were described by Ju (1990) and Martin (1969) (as
Numulariola mermillit).
SPECIMENS EXAMINED: Campeche state, Escarcega municipality, Ing.
Eduardo Sangri Serrano Forestry Experimental Station, 9.X1.88, San Martin 1180,
on wood in median subevergreen tropical rain forest (ITCV;JDR). Quintana Roo
state, Oth6n P. Blanco municipality, Tamalcab Island, 11.X1.88, San Martin 1191,
on wood in low subdeciduous tropical rain forest (ITCV;JDR); Ejido La Union,
8.XI1.86, San Martin 66, on wood in median subdeciduous tropical rain forest
(ITCV;JDR). Tamaulipas state, Ocampo municipality, El Tigre Hill, 8. VIII.88, San
Martin 990, on wood in median subdeciduous tropical rain forest (ITCV;JDR);
10. VIII.88, Chac6én 480, on wood in median subdeciduous tropical rain forest
(ITCV;JDR); GOmez Farias town 16.VII.87, San Martin 225; 14.X.88, San Martin
1122, on wood in median subdeciduous tropical rain forest (ITCV;JDR).
NOTES: This taxon poses the same problems as the previous one. We are
undecided as to whether it should be considered as a variety of Biscogniauxia
nummulana or aS a separate species (see NOTES on the previous taxon).
According to Ju (1990), this taxon and the following one cannot be distinguished
by anamorphic characters, the only apparent difference being the slightly smaller
ascospores of var. mer7illii. There is a lack of knowledge about host preferences
of both taxa.
Biscogniauxia cf. nummularia (Bull.:Fr.) Kuntze var. nummularia, Rev. Gen. Plant
2:398.. 1891.
The teleomorphis as described by Miller (1961) and Petrini & Miiller (1986)
except that Mexican material has some spores with pinched ends.
SPECIMEN EXAMINED: Chiapas state, La Trinitaria municipality, Montebello
Lagoons, 24.V.88, San Martin 802, on wood in oak-pine forest (ITCV;JDR).
NOTES: Biscogniauxia nummulania is primarily a European species. Despite the
presence of some spores with pinched ends, Mexican material is very similar to it.
On the other hand, the ascospores of this material have almost the same shape of
those of H. maculum (Schwein.) J. H. Miller, but are smaller (see Miller, 1961 for
description of H. maculum). Welden & Guzman (1978) previously reported this
species, as Hypoxylon nummularium, from the state of Veracruz.
Biscogniauxia nummularia (Bull.:Fr.) Kuntze var. "pseudopachyloma"
Hypoxylon nummularium Bull.:Fr. var. pseudopachyloma (Speg.) J. H. Miller,
Monogr. of the world species of Hypoxylon p. 125. 1961.
The teleomorph is as described by Miller (1961). The anamorph was
described by Ju (1990).
240
SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, Montes Azules
Biosphere Reserve, 27.V.88, San Martin 780, on wood in evergreen tropical rain
forest (ITCV;JDR). Nuevo Leon state, Santiago municipality, El Cercado, 23.X.88,
San Martin 1142, on wood in submontane scrub with Quercus spp. ITCV;JDR).
Tamaulipas state, GOmez Farias town, 20.VII.88, San Martin 976, on wood of
Citrus sinensis (L.) Osbeck. in an abandoned orchard.
NOTES: Material listed above seems to represent what Miller (1961) considered
to be variety pseudopachyloma of B. nummularia (as Hypoxylon). We are presently
unable to decide whether pseudopachyloma should be considered a variety of B.
nummularium or a separate species as did Martin (1969) (as Numulanrola
pseudopachyloma). (See also remarks on other putative varieties of B. nummulana
herein.)
Biscogniauxia sp. (José Castillo 1100B). Figs. 1-2, 9-11.
Stromata applanate, erumpent, covering extensive areas of the host, more
than 50 cm long x 0.4-0.6 mm thick; externally at first bluish-gray then blackish-
white, internally black. Texture hard. Surface uneven due to perithecial ostioles
and irregularities of the substrate. Perithecia ovoid to more or less pyriform, 0.2-
0.4 “m high x 0.1-0.3 mm broad. Ostioles prominently papillate surrounded by a
blackish area. Asci cylindrical, eight-spored, with uniseriate spore-arrangement,
very short-stipitate, 140-156 4m total length x 10-12 4m broad, the spore-bearing
part 132-143 zm, with apical ring bluing in Melzer’s iodine reagent, rectangular,
1.5-3 zm high x 3-4.5 4m broad. With hyaline paraphyses 8-9 4m broad at bases.
Ascospores dark brown, ellipsoid-inequilateral to navicular with narrow to rounded
ends, 18.5-20(-21) x (7-)7.5-9 4m, with straight germ slit spore-length on convex
side.
SPECIMENS EXAMINED: Tamaulipas state, Casas municipality, E] Lajeadero
Ranch, 8.X.86, José Castillo 1100B, on wood of living Quercus sp. in an oak forest
(ITCV;JDR); Km 45 Victoria City-Soto La Marina road, San Martin 1090, on
wood of Quercus sp. in an oak forest (ITCV;JDR).
NOTES: In its host preference and the whitish color of the surface in mature
specimens this fungus resembles Biscogniauxia atropunctata, but differs in the
presence of blackish areas surrounding perithecial ostioles, and smaller asci, apical
ring and ascospores (see descriptions of H. atropunctatum in Miller, 1961, and
-Biscogniauxia atropunctata herein). It might be a variant of B. atropunctata or even
a new species. We prefer to postpone this decision until cultures are established
and examined.
Biscogniauxia sp. (San Martin 725). Figs. 12-14.
Stromata applanate, erumpent, 0.9-2.5 cm long x 0.7-1 cm broad; externally
and internally black. Surface smooth with papillae. Texture carbonous. Perithecia
more or less tubular, 0.8-0.9 mm high x 0.4-0.5 mm broad. Ostioles moderately to
prominently papillate-conical. Asci cylindrical, eight-spored, with partly biseriate
241
spore-arrangement, short-stipitate, 117-135 fim total length x 6-7.5 4m broad, the
spore-bearing part 95-106 44m, with apical ring bluing faintly in Melzer’s iodine
reagent, short cylindrical to slightly cuneate, 1.5-2 4m high x 2.5-3 4m broad.
Ascospores brown, ellipsoid-inequilateral to navicular, with narrow to acute ends,
13-16(-17) x (4.5-)5-5.5(-6) Am, with germ slit straight, much less than spore-length
on the flattened side.
SPECIMEN EXAMINED: Chiapas state, San Cristobal de las Casas municipality,
E] Huitepec Hill, 4. VI.88, San Martin 725, on wood in oak forest ITCV;JDR).
NOTES: Stromatal characters of this species are reminiscent of Hypoxylon
nummularium vat. theissenit (Sydow) J. H. Miller, but the ascospores have narrow
to acute ends and the germ slit is shorter than that of the ascospores as depicted
by Miller (1961).
Camillea Fr.
Camillea Fr. includes taxa with stromata that are applanate, cylindrical, or
broadly conic-truncate. They are usually dark brown or black and highly
carbonaceous. The stroma is bipartite, the outermost layer shed to expose the
ostiolar openings. Ostioles take various forms and configurations and are highly
diagnostic for many species. A single ostiolar canal services one or several
perithecia, depending upon the species. Asci feature apical rings that blue in
Melzer’s iodine reagent, are approximately as broad as high, and are rhomboid or
diamond-shaped. Ascospores are almost hyaline, pale yellow, or light brown and
lack a germ slit. By light microscopy the ascospores appear smooth to granular or
slightly ornamented. By scanning electron microscopy they are seen to be
ornamented with warts, spines, pits, reticulations, or ribs combined with an intricate
substructure. Anamorphs where known are accommodated in Xylocladium Sydow
(= Basidiobotrys Hohnel). Many of the species now recognized as Camillea were,
until recently, included in Nummularia or Hypoxylon. A recent contribution toward
a monograph of Camillea reviews most current information on the genus (Laessge
et al., 1989).
KEY TO SPECIES OF CAMILLEA FROM MEXICO
ii Ascospores light brown to brown, verrucose or echinulate .......... Z
iL Ascospores subhyaline to light brown, punctate to reticulate or more
complexly ornamented (ornamentation seen clearly only by SEM) .... 3
fap Stromata short-cylindrical or pulvinate, protruding 3 mm or
more above the host surface level. Ostioles papillate-conical to
hemispheric. Ascospores (9-)10-11(-12) x 4-5 um .............
Bre eee. rE, «oh REI pons C. cf. verruculospora
Ze Stromata nearly applanate, or short cylindrical, not protruding
more than 3 mm above the host surface level (in our material).
Ostioles obscure to slightly papillate. Ascospores 12-16 x 5-6
242
in
11.
Level 28 eerie tat ae ones eae ac ee SEE OPC eae LL eo eS Se 4
Stromata applanate, not reaching 3 mm above the host surface level .. 6
4. Ostioles in radiating depressions. Ascospores (12-)13-15 x 3.5-
ACD EAI Gringc 22h Lora DINE, jo cl's. £48 ogee, MARIS Sale hh Feria Fees C. stellata
4, Ostioles not in radiating depressions... ee ee 5
Stromata with mucronate-labiate apex. Ascospores cylindrical, 8-9 x 4-5
PEM es ate Cra anate Capa ae atOeE ocala! cick - seaiaeaiity Gg Levies ta C. mucronata
Stromata with the upper part discoid-convex. Ascospores navicular to
somewhat crescentic, (13.5-)14-16 x 4(-4.5) um ........... C. labellum
6. Ascospores ellipsoid with one end beveled ................. 7
6. Ascospores ellipsoid-ineq uilateral, navicular, or crescentic, with
both endsmoreorlessisimilar ee Se ee ee eae 10
Ostioles in minute pits. Ascospores 8-10(-11.5) x (3.5-)4-4.5(-5) um.
Stromata chiefly on Quercus wood ................64. C. punctulata
OStIOIES NOUN MING: DIES PN ee eras esos ote cock ec suaesl ates ah eee ae 8
8. Ostioles papillate, not located in depressions. Ascospores
(7) 15-10. S(T Xx 4-8. IN oem ce cele cine l=. ences sata os C. hainesii
8. Ostioles;locatediin depressions 2) 70) 01%), .s,ersrer ea rata ere vie 9
Depressions tending to remain separate, generally 1 ostiole per depression.
Ascospores (7.5-)8-9(-10) x 4-4.5(-5) um .........-22006. C. fossulata
Depressions tending to coalesce, generally more than one ostiole per
depression. Ascospores 9-11 x 4-5 um .......... C. mexicana sp. nov.
1055 -ASCOSpores 2417 AM OL JONPET oo eres cate ie ohetsseoe Nel tare tee 11
10s." Ascospores less than 21° fim Jong 00) t.rs ok. aay degra bersteee 13
Ostioles prominently papillate, on the upper part of hemispheric projections
surrounded by depressions. Ascospores (20-)22-29(-30) x 8-9 wm ......
bite wrhay ee? Ske saat ky fl Mara ae gr LCE re a ee at ee ae Cee C. cf. gigaspora
Ostioles not on the top of hemispheric projections; stromatal surface
between ostioles planGi coche icles tales nie ye pita oe ee eles oe ant 12
12. Ascospores (19-)21-24 x (4.5-)5-6 fam. Ascus apical ring
somewhat rectangular, discrete, flaring upward, 3-4 wm high .....
AOA rch as MUN Gh UO RR ne kc ey AL as C. guzmanii sp. nov.
12. Ascospores (22-)23-33 x (7-)8-9 4m. Ascus apical ring cup-
shaped, massive, (12-)12.5-14(-14.5) 4m high x 5-6 “4m broad.....
Ss EA A, Madman AE TP 8c MOA oa aa ARS SU IE C. magnifica sp. nov.
13. Ostioles obscure to conspicuously papillate; wood beneath stromata usually
stained orange. Ascospores 13-20(-21) x 6-9 wm........... C. tinctor
13. Ostioles obscure to punctate; wood beneath stromata never orange-stained.
Ascospores (13-)15-19 x (6-)7-7.5(-8) Mm «6.2... eee eee. C. cyclisca
Camillea cyclisca (Mont.) Laessge, J. D. Rogers & Whalley, Myc. Res. 93:128.
1989.
Teleomorphic features and a Xylocladium anamorph were discussed in
Laess¢ge et al. (1989).
SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, Ejido Boca de
Chajul, VII.87, San Martin 371, on wood in evergreen tropical rain forest
(ITCV;JDR). Tamaulipas state, Gomez Farias town, VII.87, San Martin 380, on
wood in median subdeciduous tropical rain forest (ITCV;JDR); Ocampo
municipality, 16.V.88, San Martin 672, on wood in low subdeciduous tropical rain
forest (ITCV;JDR).
NOTES: This taxon seems to be related to C. finctor. The latter stains wood
beneath the stromata orange; C. cyclisca does not. The differences between these
taxa are subtle.
Camillea fossulata (Mont.) Laessge, J. D. Rogers & Whalley, Myc. Res. 93:131.
1989. Figs. 15-20.
Teleomorph is as described in Laess¢e et al. (1989) except that ascospores
are hyaline instead of dilute yellow, and ascus apical rings are diamond-shaped
_ instead of urn-shaped. Ascospore surface under SEM is punctate-reticulate.
SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, "Montes
Azules" Biosphere Reserve, 28.V.88, San Martin 811, on wood in evergreen
tropical rain forest (ITCV;JDR); Ejido Boca de Chajul, 29.V.88, San Martin 861,
on wood in evergreen tropical rain forest (ITCV;JDR).
NOTES: It appears that examination of more material with asci and ascospores
in good condition, as well as cultural data, are needed in order to achieve a better
concept of this taxon. It seems to be collected fairly regularly in the tropics, but
seldom with ascospores and asci in good condition.
Camillea cf. gigaspora (Massee) Laessge, J. D. Rogers & Whalley, Myc. Res.
33132511989.
244
Teleomorphic features are described in Miller (1961) under Hypoxylon
hemisphaericum J. H. Miller; ascospore ornamentation is as described by Rogers
(1977) for H. hemisphaericum.
SPECIMEN EXAMINED: Chiapas state, Ocosingo municipality, Ejido Pico de
Oro, 31.V.88, San Martin 884, on wood in median subdeciduous tropical rain forest
(ITCV;JDR). i
NOTES: Few ascospores were found in this material. Nevertheless, their shape
and size, and the characteristic disposition of the perithecial ostioles permitted a
certain degree of confidence about the specific identity.
Camillea guzmanii sp. nov. Figs. 28-31
Stroma fragmentarium, applanatum, erumpens, 4-5 mm longum x 3-3.5 mm
latum x 0.7-1 mm crassum, marginibus sterilibus distinctis; extus atrum; intus
nigellum. Textura dura. Perithecia ampulliformia, 0.4-0.6 mm alta x 0.3-0.4 mm
lata. Ostiola indistincta vel parum papillata. Asci octospori, cylindrici, stipitati,
188-203 4m longitudine x 7-8 4m crassi, partibus sporiferis 135-145 4m, annulo
apicali in liquore iodata Melzeri immerso cyanescente, rotundato ad basim et
sursum aliquantum infundibuliformi, 3-4 zm alto x 3-4 “wm lato. Paraphyses
persistentes, septatae, 5.5-9.5 fam latae. Ascosporae luteolae, unicellulares,
ellipsoideo-inequilaterales vel plus minusve lunatae, extremis angustis vel acutis,
sub lente laeves, per SEM observatae punctato-reticulatae, (19-)21-24 x (4.5-)5-6
fim. Rima germinationis nulla.
Stroma fragmentary, applanate, erumpent, 4-5 mm long x 3-3.5 mm broad
x 0.7-1 mm high, with distinct sterile margins; exterior black; interior dark. Texture
hard. Perithecia flask-shaped, 0.4-0.6 mm high x 0.3-0.4 mm broad. Ostioles
obscure to slightly papillate. Asci eight-spored, cylindrical, stipitate, 188-203 4m
total length x 7-8 44m broad, the spore-bearing part 135-145 fim, with apical ring
bluing in Melzer’s iodine reagent, rounded at the base and flared upward, 3-4 “4m
high x 3-4 fém broad. Paraphyses persistent, septate, 5.5-9.5 fm broad.
Ascospores pale yellow, unicellular, ellipsoid-inequilateral to somewhat crescentic,
with narrow to acute ends, smooth by light microscopy, punctate-reticulate by SEM,
Figs. 21-31. Camillea labellum and C. guzmanii. 21-27. C. labellum. 21.
Ascospore, x 2425. 22. Ascospore in upper part of ascus, x 5000. 23. Ascus
apical ring, x 2900. 24. Ascospore showing ornamentation, x 5000. 25. Detail of
ascospore ornamentation, x 15000. 26. Anamorph. Ampulla with conidiogenous
cells and conidial primordium (arrow),-x 1650. 27. Conidiogenous cells bearing
conidia (arrow), x 1650. 28-31. C. guzmanit. 28. Ascospore, x 1400. 29. Ascus
apical ring, x 2300. 30. Ascospore showing ornamentation, x 4000. 31. Detail of
ascospore ornamentation, x 15000.
Figs. 21, 22, 26-28 by DIC. Figs. 23 and 29 by BF. Figs. 24, 25, 30, 31 by
SEM. Figs. 21, 22, 26-28 from water mounts. Figs. 23 and 29 from mounts in
Melzer’s reagent.
245
LQE <
246
(19-)21-24 x (4.5-)5-6 fm. Lacking a germination slit.
SPECIMEN EXAMINED: Veracruz state, Catemaco municipality, Catemaco-
Montepio road, 4.X1.88, San Martin 1399, on wood in evergreen tropical rain forest
(ITCV,HOLOTYPE).
NOTES: Camillea guzmanii differs from Camillea magnifica described herein, in
having obscure to slightly papillate ostioles instead of punctate ones, in having a
much smaller ascus apical ring, ie. 3-4 2m high x 3-4 “wm broad vs (12-)12.5-
14(-14.5) um high x 5-6 4m broad, and in having shorter ascospores, i.e. (19-)21-24
x (4.5-)5-6 fam vs (22-)23-33 x (7-)8-9 Am. Both species have a similar poroid-
reticulate ascospore ornamentation seen by SEM, but pores in C. guzmanii are
more elongated. The Mexican fungus shares with Camillea hyalospora (Pat.) J. D.
Rogers, Laessge, & Lodge its slightly papillate ostioles, but the latter has larger
ascospores, i.e. 26-31 x (5-)7.5-9 zm. Ascospore ornamentation of C. hyalospora
as seen by SEM is poroid-reticulate but has more elongated pores than those of
C. guzmanii [see Rogers, 1975b for ascospore ornamentation of N. hyalospora (as
Hypoxylon glycyrrhiza Berk. & M. A. Curtis), and Rogers et al., 1991 for description
of C. hyalospora}.
Camillea guzmanii is named after the eminent Mexican mycologist, our
friend Gast6n Guzman.
Camillea hainesii (J. D. Rogers & Dumont) Laess¢e, J. D. Rogers & Whalley, Myc.
Res. 93:134. 1989.
Teleomorph was described by Rogers & Dumont (1979) as Hypoxylon
hainesii based on fragmented material. Stromata applanate, orbicular, erumpent,
with sterile margins, 0.9-2.5 cm long x 5-7 mm broad x 0.3-0.8 mm high. Ascospores
rectangular to ellipsoid, with one end somewhat acute, the other rounded, (7-)7.5-
10.5(-11) x 4-4.5 wm.
SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, "Montes
Azules" Biosphere Reserve, 30.V.88, San Martin 823 and 872, on wood in
evergreen tropical rain forest (ITCV;JDR). Oaxaca state, Temazcal town, 8.X.88,
San Martin 1093, on wood in median subdeciduous tropical rain forest
(ITCV;JDR).
NOTES: Compared with type material of C. hainesti (see Rogers & Dumont,
1979), the Mexican collections differ in having ascospores more variable in shape
and in being slightly larger both in length and width.
Camillea labellum Mont., Ann. Sci. nat. Bot. 4:113. 1855; Sensu Dennis, Kew Bull.
2:319. 1957. Figs. 21-27.
Teleomorph described in Dennis (1957) and Laessge et al. (1989).
Ascospores are smooth under LM, but are seen by SEM to be composed of closely
packed rod-shaped elements.
Colonies covering 9 cm diam Petri dish in 11 days, with mycelium at first
247
white with yellowish tones, then turning pale-yellow, floccose, becoming felty in old
colonies; margin indistinct, with peripheral hyphae woolly to somewhat appressed.
Odor strongly yeasty which by continuous transferring gradually disappears.
Reverse at first yellowish then greenish, finally dark-green. Conidia produced in
15 days. Hyphae septate, hyaline, of two types: one torulose ca. 4 4m broad, the
other even, 6 4m broad. Conidiophores upright, with determinate growth, arising
singly and laterally from hyphae, each composed of a mononematous, smooth stipe
more than 450 4m long, terminated by a swollen ampulla. Ampullae subglobose
to clavate, 10-12 x 16-18 um, bearing conidiogenous cells, with poroid scars at the
point of conidiogenous cell attachment. Conidiogenous cells hyaline, continuous,
oblong-elliptical, inflated at the apex, numerous, often closely compacted on the
surface of the ampullae, 5-9 4m high x 2-4 44m broad, with conspicuous secession
scars. Conidia grayish in mass, hyaline individually, borne on the upper part of
conidiogenous cells or directly from apparently unspecialized hyphae, ellipsoid to
somewhat allantoid with one end rounded, the other truncate to somewhat
attenuated, bearing basal scar indicating former point of attachment to
conidiogenous locus, 6-11(-13) x (2-)2.5-3 zm, with internal oil droplets.
SPECIMENS EXAMINED: Campeche state, Escarcega municipality, "Ing.
Eduardo Sangri Serrano" Forestry Experimental Station, 9.X1.88, San Martin
1181C, on wood in median subevergreen tropical rain forest (ITCV;JDR). Chiapas
state, Ocosingo municipality, Ejido Boca de Chajul, 29.V.88, San Martin 839, on
wood in evergreen tropical rain forest (ITCV;JDR); Chancal4, Rio Chancala, Alt
100 m., 12.23.85, Chacon-Zapata 3346, on wood in evergreen tropical rain forest
(XAL). Quintana Roo state, Oth6n P. Blanco municipality, Ejido La Union, IX.87,
San Martin 313, on wood in median subdeciduous tropical rain forest (ITCV;JDR);
San Felipe Bacalar, 10.X1.88, San Martin 1219 and 1233 (CULTURED), on wood
in an acahual (ITCV;JDR). Tamaulipas state, Ocampo municipality, 16.V.88, San
Martin 663, on wood in low subdeciduous tropical rain forest (ITCV;JDR).
Veracruz state, Catemaco municipality, "Los Tuxtlas" UNAM Biological Station,
4.X1.88, San Martin 1264, on wood in evergreen tropical rain forest ITCV;JDR).
NOTES: Mexican material matches the concept of Dennis (1957) for C. labellum.
The Xylocladium anamorph supports the hypothesis that the remaining members
of the genus Camillea for which the anamorphs have not yet been obtained, will
likewise have a Xylocladium state.
Camillea magnifica sp. nov. Figs. 3-5, 32-35
Stromata erumpentia, applanata, circularia vel irregularia, restricta, 0.2-3.4
cm longa x 0.15-1.4 cm lata x 0.7-0.8 mm alta, marginibus sterilibus aliquantum
elevatis; extus brunneola vel nigella; intus nigella. Textura dura. Perithecia
ampulliformia, 0.7 mm alta x 0.3-0.4 mm lata. Ostiola indistincta vel subtiliter
punctata vel parum papillata. Asci octospori, cylindrici, breviter stipitati, 185-210
Hm longitudine tota x 9-10 zm crassi, partibus sporiferis 157-177 4m, annulo
apicali in liquore iodata Melzeri immerso cyanescente, cupulato, magno, (12-)12.5-
14(-14.5) um alto x 5-6 zm lato. Paraphyses persistentes, septatae, longior quam
asci, 4-6 4m latae. Ascosporae luteolae, unicellulares, ellipsoideo-inequilaterales
248
vel plus minusve lunatae, extremis angustis vel acutis, sub lente subtiliter aspris, per
SEM observatae punctato-reticulatae, (22-)23-33 x (7-)8-9 Am. Rima germinationis
nulla.
Stromata erumpent, applanate, circular or irregular, restricted, 0.2-3.4 cm
long x 0.15-1.4 cm broad x 0.7-0.8 mm high, with somewhat raised sterile margins;
exterior brownish to black; interior blackish. Texture hard. Perithecia flask-
shaped, 0.7 mm high x 0.3-0.4 mm broad. Ostioles obscure to finely punctate to
slightly raised. Asci eight-spored, cylindrical, short-stipitate, 185-210 4m total
length x 9-10 4m broad, the spore-bearing part 157-177 vm, with apical ring bluing
in Melzer’s iodine reagent, cup-shaped, massive, (12-)12.5-14(-14.5) 4m high x 5-6
fm broad. Paraphyses persistent, septate, longer than asci, 4-6 (4m broad.
Ascospores yellowish, unicellular, ellipsoid-inequilateral to somewhat crescentic,
with narrow to acute ends, finely roughened by LM, punctate-reticulate by SEM,
(22-)23-33 x (7-)8-9 Am. Lacking germination slit.
SPECIMEN EXAMINED: Tamaulipas state, GOmez Farias town, no date, San
Martin 258, on wood in median subdeciduous tropical rain forest (ITCV,
HOLOTYPE; JDR;ISOTYPE).
NOTES: Camillea magnifica has ascospores in the size range of C. gigaspora
(Massee) Laess¢ge, J. D. Rogers & Whalley but its ostioles are punctate instead of
papillate on raised hemispherical elevations, and its apical ring is much larger, i.e.,
(12-)12.5-14(-14.5) 4m high vs 4.5-6.5 4m high. The ascospore surface as seen by
SEM is poroid-reticulate in both species [see ascospore ornamentation of C.
gigaspora (as Hypoxylon hemisphaericum J. H. Miller) in Rogers, 1977]. See
NOTES on Camillea guzmanii herein for differences between C. guzmanii and C.
magnifica). The name magnifica refers to the large apical ascus ring.
Camillea mexicana sp. nov. Figs. 6-8, 36-39
Stromata erumpentia, applanata, ca 1 m longa x 7-10 cm lata x 0.4-0.5 mm
alta, marginibus sterilibus; extus brunneola vel nigella; intus nigella. Textura dura.
Perithecia oblonga, 0.4-0.5 mm diam. Ostiola subliter papillata, 2-4 disposita in
depressionibus non profundis et saepe coalescentibus. Asci octospori, cylindrici,
breviter stipitati, 104-119 “zm longitudine tota x 6-6.5 1m crassi, partibus sporiferis
68-78 4m, annulo apicali in liquore iodata Melzeri immerso cyanescente, valde
Figs. 32-39. Camillea magnifica and C. mexicana. 32-25. C. magnifica. 32.
Ascospore, x 1240. 33. Ascus apical ring, x 1750. 34. Ascospore showing
ornamentation, x 3500. 35. Detail of ascospore ornamentation, x 11000. 36-39.
C. mexicana. 36. Ascospore, x 23000. 37. Ascus apical ring, x 1625. 38.
Ascospore showing ornamentation,x 8000. 39. Detail of ascospore ornamentation,
-x 15000.
Figs. 32 and 36 by DIC. Figs. 33 and 37 by BF. Figs. 34, 35, 38, 39 by SEM.
Figs. 32 and 36 from water mounts. Figs. 33 and 37 from mounts in Melzer’s
reagent.
250
rhombeo, 3 um alto x 4 fm lato. Paraphyses persistentes, septatae, 7-10 4m latae.
Ascosporae hyalinae vel luteolae, unicellulares, plus minusve oblongae, extremo
saepe devexo, sub lente laeves, per SEM observatae plus minusve punctato-
reticulatae, 9-11 x 4-5 4m. Rima germinationis nulla.
Stromata erumpent, applanate, ca. 1 m long x 7-10 cm broad x 0.4-0.5 mm
high, with sterile margins; exterior brownish to blackish; interior blackish.
Perithecia oblong, 0.4-0.5 mm diam. Ostioles finely papillate, in groups of 2-4 in
shallow depressions which frequently coalesce. Asci 8-spored, cylindrical, short-
stipitate, 104-119 zm total length x 6-6.5 44m broad, the spore-bearing part 68-78
jum, with apical ring bluing in Melzer’s iodine reagent, strongly rhomboid, 3 “4m
high x 4 4m broad. Paraphyses persistent, septate, 7-10 im broad. Ascospores
hyaline to pale yellow, unicellular, more or less rectangular, often sloping at one
end, smooth by light microscopy, more or less punctate-reticulate by SEM, 9-11 x
4-5 um. Lacking germination slit.
SPECIMEN EXAMINED: Chipas State, Ocosingo municipality, "Montes Azules"
Biosphere Reserve, 29.V.88, San Martin 856, on wood in evergreen tropical rain
forest ITCV,HOLOTYPE;JDR,ISOTYPE).
NOTES: Camillea mexicana differs from C. fossulata (Mont.) Laessge, J. D.
Rogers & Whalley in having a stromatal surface that is blackish or brownish-black
instead shiny-black, in its faint, more indefinite surface depressions which tend to
coalesce rather than remaining separate, in having several ostioles per depression,
in having slightly larger ascospores, i.e. 9-11 x 4-5 zm vs (7.5-)8-9(-10) x 4-4.5(-5)
fim, and in having unique interlocked, puzzle-like ascospore ornamentation as seen
by SEM rather than poroid-reticulate [see description of C. fossulata in Miller,
1961, (as Hypoxylon fossulatum Mont.), and herein].
Camillea mexicana approximates Camillea flosculosa (Starb.) Laessge, J. D.
Rogers & Whalley in the disposition of the ostioles and ascospore morphology seen
by LM, but differs in having smaller ascospores, i.e. 9-11 x 4-5 fam vs 11-15.5 x 5-
6.5 44m, and in having the ascospore ornamentation referred to above instead of
poroid-reticulate [see Rogers, 1977, for description of ascospore ornamentation in
Hypoxylon flosculosum (Starb.) J. H. Miller]. The Mexican fungus shares with
Camillea punctulata (Berk. & Ravenel) Laessge, J. D. Rogers & Whalley the shape
of the ascospores and to a lesser degree the ascospore ornamentation, but the
latter species has pitted ostioles instead of slightly papillate ones in faint
depressions.
This highly distinctive fungus seems to have a transitional type of ascospore
ornamentation from poroid reticulate to ribbed anastomosing, or vice versa.
Camillea mucronata Mont., Ann. Sci. Nat. Bot. 4:112. 1855.
Teleomorph is as described by Medel & Chac6n (1988), and Laess¢e et al.
(1989).
SPECIMEN EXAMINED: Chac6n-Zapata 3316, 22.XII.85, Michol-Ha falls,
Palenque municipality, Chiapas state, on wood in evergreen tropical rain forest
(XAL).
251
NOTES: Medel & Chacén (1988) reported the species from the Mexican state of
Chiapas.
It is interesting to note that, with one exception, material of C. mucronata
so far reported was collected near rivers, suggesting that the species is confined to
some tropical, riparian host(s) (see Laessge et al. 1989 for distribution of C.
mucronata).
Camillea obularia (Fr.) Laessge, J. D. Rogers & Lodge, Mycologia 83:224. 1991.
Camillea broomeiana (Berk. & M. A. Curtis) Laessge, Rogers & Whalley, Myc.
Res. 93:149. 1989.
Teleomorph is as described in Laess¢ge et al. (1989) under C. broomeiana;
the anamorph was described in Rogers (1975a) [as Nummulania broomeiana (Berk.
& M. A. Curtis) J. H. Miller].
SPECIMENS EXAMINED: Oaxaca state, Km 65 Loma Bonita-Sayula off-side
road, 21.V.88, San Martin 719, on wood of Delonix regia (Bojer ex Hook) Raf.,
("flambollan"), (ITCV;JDR). Tamaulipas state, Gomez Farias town, 17.1.85, Garcia
J. 4599, on wood of Mangifera indica L. ("mango"), in an orchard (ITCV;JDR).
NOTES: Mexican material fits well the species concept of C. obularia. It is
interesting to note that out of 3 collections made by F. C. Deighton in Sierra
Leone, one was on Delonix regia and the other on Mangifera indica (see Laessge,
et al., 1989). Mexican material was found on those hosts (see SPECIMENS
EXAMINED above).
Camillea punctulata (Berk. & Ravenel) Laess¢ge, J. D. Rogers & Whalley, Myc.
Res. 93:143. 1989.
The teleomorph was as described by Miller (1961), and the anamorph was
described by Jong & Rogers (1972) [as Hypoxylon punctulatum (Berk. & Ravenel)
Cooke in both publications].
SPECIMENS EXAMINED: Chiapas state, La Trinitaria municipality, Montebello
Lagoons, 50 km from the Guatemalan border, 24.V.88, San Martin 789, on wood
in pine-oak forest ITCV;JDR). Nuevo Le6n state, Santiago municipality, El
Cercado, VIII.1987, San Martin 395, on wood of Quercus ryzophylla Weath. in
submontane scrub with Quercus spp. (ITCV;JDR). Tamaulipas state, Gomez
Farias municipality, Malacate, 31.11.88, Baldazo 219, on wood of Quercus aff.
germana Cham. & Schlecht. in cloud forest (ITCV;JDR).
NOTES: This species seems to be confined almost exclusively to Quercus spp.
(Barnett, 1957; Jong & Rogers, 1972). Its ascospore shape and ornamentation by
SEM approximates that of Camillea mexicana described above, but the ascospores
of the latter are larger and its ostioles are not in pits [see description of ascospore
surface of H. punctulatum in Rogers (1977), and description of C. mexicana
herein].
Medel et al. (1989) reported this species [as Hypoxylon punctulatum (Berk.
& Ravenel) Cooke] from the Mexican state of Veracruz growing on wood in a
252
cloud forest.
Camillea stellata Laessge, J. D. Rogers & Whalley, Myc. Res. 93:144. 1989.
Teleomorph is as described in Laess¢e et al. (1989).
SPECIMENS EXAMINED: Chiapas state, Palenque municipality, Palenque
Ruins, [X.1987, San Martin 296, on wood in evergreen tropical rain forest
(ITCV;JDR). Quintana Roo state, Oth6n P. Blanco municipality, Ejido La Union,
8.X11.86, San Martin 37, on wood in median subdeciduous tropical rain forest
(ITCV;JDR).
NOTES: Mexican material seems to be typical of the species even though it has
few ascospores in good condition.
Camillea tinctor (Berk.) Laessge, J. D. Rogers & Whalley, Myc. Res. 93:145. 1989.
Teleomorph and anamorph are more or less as described by Laess¢e et al.
(1989).
SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, Montes Azules
Biosphere Reserve, 27.V.88, San Martin 830, on wood in evergreen tropical rain
forest (TCV;JDR); Palenque municipality, Palenque Ruins, 2.VI.88, San Martin
876, on wood in evergreen tropical rain forest (ITCV;JDR). Nuevo Le6n state,
Santiago municipality, El Cercado, 26.V.88, Moreno 221 and 225, on wood in
submontane scrub with Quercus spp. (ITCV;JDR); 23.X.88, San Martin 1146,
(same data as Moreno 221 and 225 above); 29.1.89, Moreno 267 (CULTURED),
on wood of Morus nigra L. ("mora"), in submontane scrub with Quercus spp.,
(ITCV;JDR). Tamaulipas state, G6mez Farias municipality, "El Nacimiento’,
29.V.88, San Martin 620, on wood in median subdeciduous tropical rain forest
(ITCV;JDR); Hidalgo municipality, Ejido Los Mimbres, "Los Alamos", 7.VII.87,
San Martin 200, on wood of Platanus sp., on a river bank (ITCV;JDR). Veracruz
state, Jalapa Enriquez municipality, "Francisco Javier Clavijero" Botanical Garden,
VIII.87, San Martin 620 A, on wood in cloud forest ITCV;JDR); San Andrés
Tuxtla municipality, Eyipantla falls, 5.X1.88, San Martin 1339 (CULTURED), on
wood of a riparian tree (ITCV;JDR).
NOTES: Mexican material does not always impart an orange stain to the wood
beneath stromata; perithecial size is variable, i.e., in Moreno 225 they measure 0.2-
0.3 mm broad and in San Martin 620 "A" they measure 0.6-0.7 mm; ascospore size
is variable among collections, i.e., in Moreno 267 the range is 13-17 fam while in
San Martin 1339 it is 18-20(-21) zm; ascospores of Moreno 267 readily germinated
in water and produced germ tubes at one or both ends of the propagules, even
when stromata lacked asci and thus appeared old.
Anamorphic features are variable as well, i.e., San Martin 1339 produced
conidia highly variable in size, 6-18 x (1.5-)2-3(-3.5) 4m, and Moreno 267 produced
ampullae not as broad as in "typical" isolates, and Nodulisporium-type conidio-
phores were more common than in other isolates. It appears necessary to obtain
additional fresh collections of this species and study the teleomorphic and
253
anamorphic features in order to assess its limits.
Camillea cf. verruculospora J. D. Rogers, Laessge & Lodge, Mycologia 83:224.
1991;
The teleomorph is as described by Rogers et al. (1991), except that
ascospores of Mexican material are smaller, i.e., (9-)10-11(-12) x 4-5 zm vs (11-)12-
13.5 x 4.5-5(-6.5) wm.
SPECIMEN EXAMINED: Chiapas state, Ocosingo municipality, Ejido Boca de
Chajul, 29.V.88, San Martin 837, on wood in an acahual (ITCV;JDR).
NOTES: The Mexican collection is probably not fully mature. This might account
for the spores being smaller than usually encountered.
ACKNOWLEDGMENTS
PPNS No. 0121, Department of Plant Pathology, Project 1767, Washington
State University, College of Agriculture and Home Economics. This study was
supported by National Science Foundation Grant BSR-9017920 to JDR. We thank
the Consejo del Sistema Nacional de Educaci6n Tecnologica for furnishing support
to SM during 1986-88 for the project Ecologia y Distribuci6n de Hongos
Destructores de Madera en el Noreste de Mexico and to Consejo Nacional de
Ciencia y Tecnologia de Mexico (CONACYT) for a grant to SM during his
graduate studies. We thank Donald P. Rogers, Auburn, WA, for correcting the
Latin descriptions. We thank the following associates at Washington State
University: Y.-M. Ju for his aid in numerous taxonomic matters; Michael J. Adams
for aid with SEM and photography; Jane Lawford for reconstructing the
manuscript; Lori M. Carris for reading the manuscript. We thank Brenda E.
Callan, Forestry Canada, Victoria, B.C., Canada for reading the manuscript. We
thank the curators of cited herbaria for the loan of specimens.
SM had extensive aid and encouragement in the portion of the studies
conducted in Mexico. He thanks the following persons: Professor José Castillo
Tovar for the first lectures on mycology and encouragement; Dr. Gaston Guzman
for his friendship and encouragement; Biol. Lucrecia Garcia Alanis, Biol. Arnulfo
Moreno, Dr. Efrén Cazares Gonzalez, Biol. Santiago Chac6n Jiménez, Biol. Bertha
Baldazo, Biol. Pablo Lavin, Biol. Catalina Flores, M. en C. Gonzalo Guevara, Biol.
Jess Garcia, and Biol. Maria Concepci6n Herrera for aid in collecting and the
latter also for her aid in identifying plant material.
SM likewise thanks Biol. Santiago Chac6n-Jiménez and Biol. Arnulfo
Moreno for sharing their interest in the adventure of collecting; Dr. Alberto
Rodriguez Fernandez and Biol. Javier Chavelas Polito for providing collecting
facilities in Quintana Roo and Biol. Francisco Quinto Adrian for his friendship and
aid in Quintana Roo; Biol. Antonio Espinoza Banda for collecting facilities in
Campeche and Biol. Jorge Ayala Guajardo for his friendly aid while collecting in
the beautiful and interesting state of Chiapas.
254
LITERATURE CITED
Barnett, H. L. 1957. Hypoxylon punctulatum and its conidial stage on dead oak
trees and in culture. Mycologia 49:588-595.
Callan, B. E. and J. D. Rogers. 1990. Teleomorph-anamorph connections and
correlations in some Xylaria species. Mycotaxon 36:343-369.
Dennis, R. W. G. 1957. Further notes on tropical American Xylariaceae. Kew.
Bull. 1957. 297-332.
Glawe, D. A. and J. D. Rogers. 1986. Conidial states of some species of
Diatrypaceae and Xylariaceae. Can. J. Bot. 64:1493-1498.
Holmgren, R. K., N. H. Holmgren, and L. C. Barnett. 1990. Index Herbariorum.
Part I: The herbaria of the world. Ed. 8. New York Botanical Garden.
693 p.
INEGI. 1983. Sintesis Geografica y Nomenclators de los estados de Coahuila,
Morelos, Nuevo Leé6n y Tamaulipas. Secretaria de Programacién y
Presupuesto. México.
INEGI. 1988. Carta Topografica de México 1:250 000. Secretaria de
ProgramaciOn y Presupuesto. México.
Jong, S. C., and J. D. Rogers. 1972. Illustrations and descriptions of conidial
states of some Hypoxylon species. Wash. State Agric. Exp. Sta. Bull. 71. 51
p.
Ju, Y.-M. 1990. Studies of Xylariaceae from Taiwan. M. Sc. Thesis. Washington
State University, Pullman, WA. 171 p.
Koeppen, W. 1948. Climatologia. Fondo de Cultura Econémica. México, D.F.
478 p.
Laessge, T., J. D. Rogers, and A. J. S. Whalley. 1989. Camillea, Jongiella and
light-spored species of Hypoxylon. Myc. Res. 93:121-155.
Martin, P. 1969. Studies in the Xylariaceae: VI. Daldinia, Numulanola and their
allies. J. S. African Bot. 35:267-320.
Medel, R. and S. Chacon. 1988. Ascomycetes lignicolas de México, II. Algunos
Pyrenomycetes y Discomycetes. Micologia Neotropical Aplicada 1:87-96.
Medel, R., S. Chac6n, and G. Guzman. 1989. Especies conocidas y nuevos
registros de Hypoxylon (Sphaeriales, Xylariaceae) en México. Rev. Mex.
Mic. 5:149-168.
Miller, J. H. 1961. A monograph of the world species of Hypoxylon. Univ.
Georgia Press, Athens. 158 p.
Pérez-Silva, E. 1983. Distribucién de algunas especies del género Hypoxylon
(Pyrenomycetes) en México. An. Inst. Biol. Univ. Nac. Aut. Méx. 54:1-22.
Petrini, L. E. and E. Miller. 1986. Haupt- und nebenfruchtformen europaischer
Hypoxylon-Arten (Xylariaceae, Sphaeriales) und verwandter pilze.
Mycologia Helv. 1:501-627.
Pouzar, Z. 1979. Notes on taxonomy and nomenclature of Nummulana
(Pyrenomycetes). Ceska Mykologie. 33:207-219.
Rogers, J. D. 1975a. Nummularia broomeiana: conidial state and taxonomic
aspects. Am. J. Bot. 62:761-764.
Rogers, J.D. 1975b. The ascospore of Hypoxylon glycyrrhiza. Mycologia 67:657-
662.
255
Rogers, J. D. 1977. Surface features of the light-colored ascospores of some
applanate Hypoxylon species. Can. J. Bot. 55:2394-2398.
Rogers, J. D. and K. P. Dumont. 1979. Los hongos de Colombia VI. Two new
applanate species of Hypoxylon. Mycologia 71:807-810.
Rogers, J. D., T. Laessge and J. Lodge. 1991. Camillea: New combinations and
a new species. Mycologia 83:224-227.
Rzedowski, J. 1978. La vegetacion de México. Ed. LIMUSA. México. 431 p.
San Martin, F. and J. D. Rogers. 1989. A preliminary account of Xylaria of
México. Mycotaxon 34:283-373.
Welden, A. L. and G. Guzman. 1978. Lista preliminar de los hongos, liquenes y
myxomycetos de las regiones de Uxpanapam, Coatzacoalcos, Los Tuxtlas,
Papaloapan y Xalapa (Parte de los estados de Veracruz y Oaxaca). Bol.
Soc. Mex. Mic. 12:59-102.
APPENDIX
Location of Collecting Sites
For the exact location of the sites where the field work was conducted, the
Atlas de México and the Geographical Syntheses and Nomenclators of the
pertinent states, edited by the Instituto Nacional de Geografia y Estadistica
(INEGI, 1983, 1988) were consulted. The main locations are listed below.
A. Campeche state:
1) Escarcega municipality, Ing. Eduardo Sangri Serrano Forestry
Experimental Station, 18° 40’ Lat; 90° 45’ Long; Topographic Map
E15-6.
B. — Chiapas state:
1) Ejido Boca de Chajul, 16° 07’ Lat; 99° 52’ Long; Topographic Map
5 12;D 153;
2) Ejido Pico de Oro, 16° 20’ Lat; 90° 45’ Long; Topographic Map
F152;D153.
3) SEI wHuitepeci Hill.) 16; 645°) Lats.92° 4.43" sLongah27509m s Alt
Topographic Map E1511.
4) El Triunfo Hill, 15° 40’ Lat; 112° 48” Long; 2450 m Alt; Topographic
Map D15-2.
5) | Malpaso Dam, 16° 48’ Lat; 93° 20’ Long; Topographic Map E-15-11.
6) Montebello Lagoons, 16° 08’ Lat; 91° 42’ Long; Topographic Map
Bi512;D 153:
7) Palenque Ruins, 17° 30’ Lat; 92° 01’ Long; Topographic Map E-15-
}
256
Coahuila state:
1) | Arteaga Municipality, San Antonio de las Alazanas, 25° 16’ Lat; 10°
34’ Long; Topographic Map G14-C35.
Federal District:
1) Km 43 Mexico-Cuernavaca Road, 19° 02’ Lat; 99° 12’ Long;
Topographic Map E-14-2.
Jalisco state:
1) | Cacoma Mountains, 19° 58’ Lat; 104° 32’ Long; Topographic Map
E-13-2-5.
2) Guadalajara City, 20° 39’ Lat; 103° 18’ Long; 1550 m Alt;
Topographic Map F-13-12.
3) La Manzanilla, 20° 00’ Lat; 103° 09’ Long; 2050 m Alt; Topographic
Map F13-12.
4) Manantl4n Biosphere Reserve, 19° 37’ Lat; 104° 13’ Long;
Topographic Map E-13-2-5.
5) | Mazamitla, 19° 55’ Lat; 103° 01’ Long; Topographic Map E-13-3.
6) Tecalitl4n, 19° 28’ Lat; 103° 18’ Long; 1140 m Alt, Topographic Map
E13-3.
Morelos state:
1) La Pera Curve, 25° 14’ Lat; 99° 14’ Long; Topographic Map E14-
AS9.
Nuevo Le6n state:
1) El Chipinque, 25° 55’ Lat; 100° 22’ Long; Topographic Map G-14-
C15.
2) El Cercado, 25° 24’ Lat; 100° 08’ Long; Topographic Map G14-C36.
3) El Salto, La Ciénega, 25° 22’ Lat; 100° 14’ Long; Topographic Map
G14-C36.
4) Guadalupe City, 25° 40’ Lat; 100° 15’ Long; Topographic Map G14-
C26.
5) La Encantada, 23° 49” Lat; 99° 41’ Long; Topographic Map F14-
Al7.
6) San Roque, 25° 36’ Lat; 100° 09’ Long; Topographic Map G14-C26.
Oaxaca:
1) | Temazcal, 18° 15’ Lat; 96° 21’ Long; Topographic Map E14-6.
2) Teotitlan del Camino, 18° 08’ Lat; 96° 53’ Long; Topographic Map
E14-6.
2544
Quintana Roo state:
1)
2)
3)
4)
5)
Ejido La Uni6n, 17° 55? Lat; 887 52° Long; Topographic Map E16-4-
is
Km 12 Cafetal-Mahahual Road, 18° 47 Lat; 87° 45’ Long;
Topographic Map E16-2-5.
La Noria Ranch, 19° 40’ Lat; 88° 44’ Long; Topographic Map E16-1.
San Felipe Bacalar, 18° 40’ Lat; 88° 25’ Long; Topographic Map
E16-4-7.
Tamalcab Island, 18° 35’ Lat; 88° 13’ Long; Topographic Map E16-
4-7.
San Luis Potosi state:
1)
Tamasopo Bath Resort, 21° 50’ Lat; 99° 25’ Long; Topographic Map
F14-8.
Tabasco state:
1)
2)
Colegio Superior de Agricultura Tropical, 17° 59’ Lat; 93° 35’ Long;
Topographic Map E15-8.
La Venta Archaeological Zone, 18° 05’ Lat; 94° 03’ Long;
Topographic Map E15-1-4.
Tamaulipas state:
11)
Alta Cima, 23° 03’ Lat; 99° 12’ Long; Topographic Map F14-A49.
Barra del Tordo, 23° 03’ Lat; 97° 45’ Long; Topographic Map F14-
B44.
Ejido Conrado Castillo, 23° 57’ Lat; 99° 28’ Long; Topographic Map
F14-A18.
Ejido Julilo, 23° 08’ Lat; 99° 12’ Long; Topographic Map F14-A49.
EI Bernal Hill, 22° 45’ Lat; 98° 35’ Long; Topographic Map F14-
B52.
El Cielo Ranch, 23° 05’ Lat; 99° 11’ Long; Topographic Map F14-
A49.
El Madrojfio, 23° 36’ Lat, 99° 13’ Long; Topographic Map F14-A29.
El Tigre Hill, 24° 37° Lat; 98° 37° Long; Topographic Map G14-D62.
Gomez Farias town, 23° 02’ Lat; 99° 09’ Long: Topographic Map
F14-A49.
Km 2 Victoria City-Matamoros City Road, 22° 02’ Lat; 99° 08’ Long;
Topographic Map F14-A19.
La Libertad, 23° 47’ Lat; 99° 11’ Long; Topographic Map F14-A19.
Veracruz State:
1)
Axocoapan, 19° 12’ Lat; 97° 00’ Long; Topographic Map E14-3.
258
2)
3)
4)
Eyipantla Falls, 18° 25’ Lat; 95° 13’ Long; Topographic Map E15-1-
4.
Jalapa, 19° 32’ Lat; 96° 55’ Long; 1440 m Alt; Topographic Map
E14-3.
Los Tuxtlas UNAM Biological Station, 18° 33’ Lat, 95° 04’ Long;
Topographic Map E15-1-4.
MY COTAXON
Volume XLVII, pp. 259-274 April-June 1993
NEW FUNGI FROM YUCCA: PLANISTROMELLA YUCCIFOLIORUM,
GEN. ET SP. NOV., ITS ANAMORPH, KELLERMANIA YUCCIFOLIORUM,
SP. NOV., AND PLANISTROMELLA UNISEPTATA, SP. NOV.,
THE TELEOMORPH OF KELLERMANIA YUCCIGENA
ANNETTE W. RAMALEY
7 Animas Place
Durango, Colorado 81301
Planistromella, a new genus closely related to Planistroma, is described. The
anamorph of Planistromella yuccifoliorum is Kellermania yuccifoliorum, a new
species. The anamorph of Planistromella uniseptata is Kellermania yuccigena.
Key words: Loculoascomycete, Coelomycete, Planistromella, Kellermania.
INTRODUCTION
Stagonospora gigantea Heald & Wolf (3) was described from Agave americana
Linnaeus, and has been reported from several species of Yucca in the United
States (2). The name has been a catch-all designation for coelomycetes from
Yucca which have unappendaged, large, hyaline, septate conidia born singly on
conidiogenous cells lining a thick-walled pycnidial chamber, because there were
no other satisfactory described genera for their disposition. The type of S.
gigantea was examined (10), and Piptarthron uniloculare Ramaley was estab-
lished to accommodate some of the fungi formerly called S. gigantea. Further
study and collections indicate that another anamorph species can be reliably
separated from the fungi placed in S. gigantea for lack of an alternative taxon.
This new anamorph, Kellermania yuccifoliorum Ramaley, has finger-shaped, 2-
septate, somewhat pointed conidia that often have a small apical appendage.
The teleomorphs of K. yuccifoltorum and K. yuccigena Ellis & Everhart
comprise a new ascomycete genus, Planistromella, which has septate ascospores
but otherwise resembles Planistroma, the teleomorph for the Pzptarthron species
in which a teleomorph has been observed (9, 10, and unpublished).
MATERIAL and METHODS
Lactophenol-cotton blue was utilized when necessary in the study of appendage
formation during conidiogenesis.
260
Cultures were obtained by suspending freshly extruded conidia in a drop of
sterile tap water and spreading the drop over the surface of commercial potato
dextrose agar (PDA) in a petri plate. Germinating conidia were isolated onto
plates of fresh PDA and grown at room temperature in incident light. Asci
were washed 3 times in sterile tap water and placed on petri plates of PDA.
Growth was isolated onto petri plates of fresh PDA and grown at room
temperature in incident light.
Planistromella yuccifoliorum Ramaley and Kellermania yucctfoliorum Ramaley
were discovered among specimens in a loan from the University of California at
Berkeley (UC). Fresh K. yucctfoliorum was collected subsequently from the
desert of the southwestern United States at eleven sites from Washington
County, Utah, southward to San Bernardino and Riverside Counties, California,
and Cocconino County, Arizona, on Yucca baccata Torrey, Y. brevifolia
Engelmann, and Y. thornberi McKelvey. |
Measurements were made of fungal tissue mounted in sterile tap water. Ten
conidia were measured from one conidioma on each of five leaves in the eleven
fresh collections. Material in two of the herbarium specimens was very limited,
so only twenty-five conidia for each of those two specimens (UC 1138875, UC
1202973) were measured. An analysis of variance (ANOVA) of conidial width
for all groups was prepared. The sum of squares simultaneous test procedure
(SS-STP) was carried out for: (a) all collections, (b) fresh vs herbarium
collections, (c) all fresh collections, (d) fresh collections from Y. brevifolia vs
those from other Yucca species, and (e) fresh collections from Y. brevifolia only
(Table 1) to test the significance of differences in mean conidial width for these
groups.
TAXONOMY
PLANISTROMELLA Ramaley, gen. nov.
Stromata atrocolorata, subepidermalia, immersa, erumpentia, separata vel
confluentia, glabra; paries crassus, cupulatus, pars supera in epidermide hospitis
affixa, ex aliquot stratts cellularum atroparietum compositus. Stratum
supremum stromatis ex columnis cellularum elongatarum compositum. Loculi
ascogeni in eodem stromate ac loculi conidiogent vel spermatiogeni vel ambo
ezortentes. Locult ovoidet vel globost, unistrati, textura interthecio repletz,
omnes ostiolati periphysibus, per columnas ex cellulis elongatis compositis
separatt. Asci bitunicati, subclavatt vel prope cylindrati. Ascosporae leves,
hyalinae, septatae.
Typus generis: Planistromella yuccifoliorum Ramaley
Stromata dark, subepidermal, immersed, erumpent, separate or confluent,
glabrous; wall thick, cup-shaped, upper portion attached to the host epidermis,
composed of several layers of dark-walled cells. The top of a stroma composed
261
of columns of elongated cells. Ascogenous locules developing in the same
stroma as the conidial or spermatial locules or both. Locules ovoid to globose,
in a single layer, filled with abundant interthecial tissue, each with a
periphysate ostiole, separated by columns of elongated cells. Asci bitunicate,
slightly clavate or nearly cylindric. Ascospores smooth, hyaline, septate.
Planistromella yuccifoliorum Ramaley sp. nov.----- Big el ee UA wks he
Stromata fusca vel atra, subepidermalia, tmmersa, erumpentia, amphigena,
separata vel confluentia, glabra, 300-550 pm diam ubi separata, usque ad 450
pm elata, 1-5 locularia; paries crassus, cupulatus, pars supera in epidermide
hospitis affiza, ex 6-12 stratis cellularum atroparietum compositus. Stratum
supremum stromatis ex columnis cellularum elongatarum compositum. Locult
ascogeni in eodem stromate ac locult conidiogeni vel spermatiogenit vel ambo
exorientes, asct post conidia vel spermatia vel ambo formatt. Loculi ovorder vel
globosi, untstrati, omnes ostiolati pertphystbus, per columnas ex cellulis
brunneis, elongatis, aliquantum crassiparietibus compostitis separati. Asci
bitunicati, subclavati, octosport, 120 x 32 pm ante prolationem ultimam et
misstonem ascosporarum. Ascosporae hyalinae, ellipsoideae aspectu uno,
curvatae ertremis obtuse rotundatis ubt 90 gradibus versae, 2-septatae, 34-42 x
10-16 pm. Textura intertheciu abunda, filamentosa, copiosa ubi asci vetissimi
maturi, fatiscens ab apice locult versus imum.
Stromata (Fig. 2, A) dark brown to black, subepidermal, immersed, erumpent,
amphigenous, separate or confluent, glabrous 300-550 ym diam when separate,
up to 450 pm tall, 1-5 loculate; wall thick, cup-shaped, the upper part attached
to the host epidermis, composed of 6-12 layers of dark-walled cells. The top of
a stroma composed of columns of elongated cells. | Ascogenous locules
developing in the same stroma as a conidiogenous or spermatiogenous locule or
both, asci formed after the production of conidia or spermatia or both. Locules
(Fig. 1) ovoid to globose, in a single layer, each with a periphysate ostiole,
separated by columns of brown, rather thick-walled, elongated cells. Asci (Fig.
2, C) bitunicate, slightly clavate, 8-spored, 120 x 32 ym before the final
elongation and ascospore release. Ascospores (Fig. 2, C) hyaline, ellipsoid in
one view, curved with bluntly rounded ends when rotated 90 degrees, 2-septate,
34-42 x 10-16 ym. Interthecial tissue abundant, filamentous, present in large
quantity when the oldest asci are mature, breaking down at the top of the locule
and proceeding downward.
Substrate: Leaves of Yucca species.
HOLOTYPE: USA. CALIFORNIA: San Bernardino County, Roadside 20
miles east of Baker (Hwy. 91/466), on leaves of Yucca brevifolia Engelmann, 14
April 1960, Isabelle Tavares No. 466, UC 12029738.
The collection of this fungus was more than thirty years old, so cultivation was
262
mes,
4 ?
Gaffe,
: He
OA
PIECE L AUP IML ISS PRK
Tei ASN
Dg
SANG
Wh I
VEE OT OSREE
Fig. 1. Planistromella yuccifoliorum, detail of an ascoma. Standard line = 50
pm.
not possible. Only dead tissues were available for measurement. In the living
state, the ascospores and asci could be expected to be larger than the above
measurements--up to 18% longer and 25% wider for the ascospores, and up to
20% longer and 26% wider for the asci (1).
Anamorph: Kellermania yuccifoliorum Ramaley sp. nov.----- Fig. 3,A-D,
Fig. 4, A & B.
263
Conidiomata amphigena, subepidermalia, atra, immersa, erumpentia etiam
obtecta in margine per epidermidem, separata vel confluentia, glabra, 250-600
(-800) xm diam ubi separata, usque ad 600 ym elata, unilocularia, ostiolata;
parties crassus, cupulatus, in eptdermide hospitis affizus, er 6-12 stratts
cellularum atrorum crassiparietum compositus, pallidior versus partem
interiorem, et 2-4 stratis cellularum hyalinarum. Stratum supremum contdio-
matis ex columnis cellularum aurearum elongatarum compositum. Ostiolum
cellulis atripartetibus cinctum, per rumpentem vel disrumpentem epidermidis
hospitts in margine interiore parietis cupulati expositum. Ontogenesis
conidiorum holoblastica. Conidiophora carentia. Cellulae macroconidiogenae
in parietibus lateralibus et basalibus conidiomatis formatae, brevicylindratae,
hyalinae, leves, omnes contdium unum acrogenum formantes. Macroconidia
anguste ellipsoidea-cylindrata, extremis basalibus obtuse rotundatis, apicibus
acutis et appendicibus usque ad 5 pm longis superpositis, plerumque 2-septata,
50-100 x (8-)13-14(-16) pm. Cellulae microconidiogenae in pariete supero
conidiomatis et canale ostiolt erorientes. Microconidia acrogena, plus minusve
cylindrata, septata, levia, hyalina, 5-10 x 2.5-4 pm. Spermatia in loculo
centrali stromatis, cum vel sine conidits, vel in uno vel plus loculis in columnis
verticalibus abundantibus cellularum elongatarum ad partetes laterales aliquot
conidiomatum affixis formata, omnis loculus ostiolatus vel locult coalescentes et
ostiolum solitarium formantes. Cellulae spermatiogenae discretae vel integratae
in conidiophoro unicellulario, cylindratae vel elongatae-conicae, phialidicae, 8-
16 r 2-3.5 pm. Spermatia bacillaria, hyalina, aseptata, levia, 3-7 © 1.6-2.5 pm.
Conidiomata (Fig. 3, A) amphigenous, subepidermal, dark, immersed, erumpent
but a rim remaining covered by the epidermis, separate or confluent, glabrous
250-600 (-800) zm diam when separate, up to 600 ym tall, unilocular, ostiolate;
wall thick, cup-shaped, attached to the host epidermis, composed of 6-12 layers
of dark, thick-walled cells, lighter toward the interior, and 2-4 layers of hyaline
cells. The top layer of a conidioma composed of vertical columns of golden
elongated cells. Ostiole surrounded by dark-walled cells, exposed by rupture or
breaking away of the host epidermis at the inner edge of the cup-shaped wall.
Conidiogenesis holoblastic. Conidiophores absent. Macroconidiogenous cells
(Figs. 3, A; 4, A) formed on lateral and basal walls of a conidioma, short-
cylindric, hyaline, smooth, each forming a single acrogenous conidium.
Macroconidia (Fig. 4, B) narrowly ellipsoid-cylindric, the base bluntly rounded,
the apex more pointed and often surmounted by an appendage up to 5 pm long,
mostly 2-septate, 50-100 x (8-)13-14(-16) wm. Microconidiogenous cells arising
on the upper wall of a conidioma and in the ostiolar channel. Microconidia
acrogenous, more or less cylindric, aseptate, smooth-walled, hyaline 5-10 x 2.5-4
pm (Fig. 3, B). Spermatia formed in the central locule of a stroma, with
conidia or without, or in one or more locules in the abundant vertical columns
of elongated cells attached to the lateral walls of some conidiomata (Fig. 3, D),
each locule ostiolate or coalescing and forming one ostiole. Spermatiogenous
cells discrete or integrated on a one-celled (usually) conidiophore (Fig. 3, C)
cylindric to elongate-conical, phialidic, 8-16 x 2-3.5 ym. Spermatia bacillar,
hyaline, smooth, 3-7 x 1.5-2.5 ym (Fig. 3, C).
264
Fig. 2. A, C, Planistromella yucctfoliorum. A. Diagram of stroma with two
ascomata sectioned. The left ascoma is drawn in detail in Figure 1; a=ascoma,
e=host epidermis, f=columnar filamentous portion of stroma, u=collapsed
locule which contained conidia or spermatia or both, w=wall. C. Asci and
ascospores. B. Planistromella uniseptata, asci and ascospores. Standard line =
A, 175 pm; B, C, 25 pm.
265
Substrate: Leaves of Yucca species.
HOLOTYPE: USA. ARIZONA: Mohave County, dirt road 0.3 miles from
mile 20, Gem Acres Road turnoff from Highway 40, on leaves of Yucca
brevifolia Engelmann, 15 April 1992, Annette Ramaley No. 9208, UC 1475101.
Paratypes: (1) USA. CALIFORNIA: Los Angeles County, Claremont,
Rancho Santa Ana Botanic Garden, leaves of Yucca baccata Torrey, 26 March
1957, Lee Bonar, UC 1138875; (2) USA. CALIFORNIA: San Bernardino
County, roadside 20 miles east of Baker (Highway 91/466), on leaves of Yucca
brevifolia Engelmann, Isabelle Tavares No. 466, 14 April 1960, UC 1202973; (3)
USA. CALIFORNIA: Riverside County, near Cottonwood Springs, Joshua
Tree National Monument, on Yucca schidigera Roezl ex Ortgies, 6 April 1952,
Lee Bonar, UC 966401; (4) USA. ARIZONA: Santa Cruz County, mile 282.8
along Interstate Highway 10, leaves of Yucca thornbert McKelvey, 13 April
1992, Annette Ramaley No. 9211, UC 1475100; (5) USA. ARIZONA: Santa
Cruz County, mile 283.6 along Interstate Highway 10, leaves of Yucca thornberi
McKelvey, 13 April 1992, Annette Ramaley No. 9212, BPI 1112456; (6) USA.
UTAH: Washington County, Visitor’s Center, just over border from Arizona
along Interstate 15, on leaves of Yucca brevifolia Engelmann, 15 April 1992,
Annette Ramaley No. 9207, BPI 1112455.
Kellermania yuccifoliorum is distinguished from the other Kellermania species
by conidial septation and the size of the apical appendage. K. anomala (Cooke)
Hohnel (Fig. 4, C) and K. yuccigena (Fig. 4, E) conidia have long apical
appendages (12-32 ym). However, appendages of K. anomala are often difficult
to observe without phase optics, and may be absent entirely. They seem pliant,
and are sometimes free from the conidia. K. yuccifoliorum conidia (Fig. 4, B)
have a short apical appendage (up to 5 ym) when an apical appendage is
present. K. yuccigena conidia regularly have one, approximately median
septum. K. anomala conidia have two septa, and the middle cell is regularly
shorter than either end cell. K. yuccifoliorum conidia have two septa, but the
middle cell is generally longer than the end cells. However, K. anomala and K.
yuccifoliorum conidia lacking an appendage may be indistinguishable if the
middle cell length is atypical for the species (compare conidia in Fig. 4, B & C).
Conidia from the fresh collections germinated readily and formed colonies with
white aerial, surface, and subsurface mycelium. The reverse of the colonies from
some collections was pale pink, from others, white. The majority of the
mycelium of colonies of most isolates soon became gray to nearly black. Within
a month, conidiomata were formed that produced typical K. yucctfoltorum
conidia. Planistromella yuccifoliorum could not be cultivated to confirm its
connection with K. yuccifoliorum. The connection is circumstantial but
assumed because (a) closely related taxa (9, 10) have similar teleomorph and
anamorph phases; (b) many collections have been made throughout the range
of the host, and the unilocular representatives of the Kellermania-Piptarthron
complex now known from Yucca brevifolia are P. uniloculare, K. yuccifoliorum,
and K. anomala. The teleomorph for P. uniloculare, Planistroma obtusilunatum
266
Ramaley, has been confirmed experimentally (10). Two stromata with a perfect
state present were found in a collection in which K. anomala was the only
unilocular Kellermania-Piptarthron representative . A single ascus with mature
ascospores was available. The ascospores were approximately medianly 1-
septate; (c) K. yuccifoliorum was the only unilocular representative of the
Kellermania-Piptarthron complex present in the collection in which P.
yuccifoltorum was found.
The average size of fifty conidia from the eleven fresh collections ranged from
66.7-80.1 x 12.4-14.2, X=72.1 x 13.2 wm. Conidia from the herbarium material
averaged 61.4 x 10.2 wm, smaller than any of the averages for the fresh conidia.
The ANOVA showed extremely significant variation among the fourteen
collections (Fs=41.59, a<.001). Results of the SS-STP test for the significance
of the contributions to the sum of squares among the groups (SSgroups) by
sample means of particular interest are shown in Table 1. The critical value for
the sum of squares (CVSS) was computed conservatively using df=13, 120.
Even so, SS groups (a), (b), (c), and (e) far exceeded the CVSS, so a<<.001 ‘for
these groups.
Table 1. Sum of Squares Simultaneous Test Procedure Results for ANOVA of
the Width of Conidia from Fresh Collections and Herbarium Collections
of Kellermania yucctfoliorum.
Groups Analyzed SSgroups
(a) all collections 13/636 1515.6262*««
(b) fresh vs herbarium collections 1/648 1206.6935*««
(c) fresh collections only 10/539 = 275.9527 **x
(d) fresh: Y. brevifolia vs other sp. 1/548 88.8552**
(e) fresh: Y. brevifolia only 1/392 VIIA 33 1oeee
*#*, A=.001; **, a=.01
These results show the significant contributions to the total variation by (1) the
fresh vs herbarium source of the conidia [(a) and (b)], (2) the different species
on which the fungus was found [(c) and (d)], and (3) the plants or site from
which a collection was made--the fresh collections from Y. brevifolia were
significantly different (e).
Conidia from the herbarium collections averaged 14.8% shorter and 22.7%
narrower than those from fresh collections, and the walls appeared fairly thick,
rather than thin as in the fresh material. Both these characteristics agree with
observations from living versus dead ascomycetes (1), and percent difference in
length and width of living versus dead conidia is within the range of the percent
difference for such measurements in ascospores. These differences account for
267
the extremely large SSgroups for conidia from fresh versus herbarium material
[Table 1, (b)], and emphasize the need for caution when drawing taxonomic
conclusions from such measurements, at least for taxa with large, thin-walled
conidia.
Planistromella uniseptata Ramaley, sp. nov.-----Fig. 2, B.
Planistromellae yucctfoliorum Ramaley similis sed stromata 250-400 pm diam
ubi separata, usque ad 350 pm elata, 1-4 locularia. Asct 105-120 rt 28-36 pm
ante prolationem ultimam et misstonem ascosporarum. Ascosporae hyalinae,
eztrema obtuse rotundata vel extremum inferius plerumque aliquantum acutum,
1-septatae, 32-48 x 13.5-17.5 pm, cellula inferior 1-1.4x longitudo cellulae
superioris. Ubi asct vetissimt maturi, teztura intertheci irregulariter
filamentosa, columnae_ dissoctantes et consortiones 2-3-cellulares vel
unicellulares formantes. Cellulae irregulariformes, cylindratae, ellipsoidiae,
ovatae, vel globosae, plerumque 10-20 xt 5-8 pum.
Like Planistromella yuccifoliorum Ramaley, but the stromata 250-400 um diam
when separate, up to 350 ym tall, 1-4 loculate. Asci (Fig. 2, B) 105-120 x 28-36
pm before the final elongation and release of the ascospores. Ascospores (Fig. 2,
B) hyaline, the ends bluntly rounded or the lower end more often slightly
pointed, l-septate, 32-48 x 13.5-17.5 ym, the bottom cell 1-1.4x the length of
the top cell. When the oldest asci are mature, interthecial tissue irregularly
filamentous, the columns dissociating into 2-3-celled associations or into
individual cells which are irregularly shaped, cylindric, ellipsoid, ovate, or
spherical, mostly 10-20 x 5-8 pm.
Substrate: Leaves of Yucca elata Engelmann.
HOLOTYPE: USA. NEW MEXICO: Soccoro County, roadside, west side of
Highway 25, mile 105.4, on leaves of Yucca elata Engelmann, 12 April 1992,
Annette Ramaley No. 9217, UC 1475102.
Planistromella uniseptata was found in small amounts on leaves of Yucca elata
Engelmann. The anamorph of the fungus is widely distributed in the United
States on Yucca angustifolia Pursh (6), Y. filamentosa Linnaeus (6), Y. glauca
Nuttall (6), Y. harrtimaniae Trelease (personal observation), and Y. whipplet
Torrey (6). When a systematic search is carried out, P. uniseptata may be
expected on these species as well as Y. elata.
Colonies arising from asci on PDA had white aerial, surface, and subsurface
mycelium with a pink reverse. After 6 weeks, conidiomata which contained
typical Kellermania yuccigena conidia had formed.
Anamorph: Kellermania yuccigena Ellis & Everhart---Fig. 4, D & E.
268
Kellermania yuccigena has been illustrated and described in modern terms (6,
12). A few additions must be made for a more complete description, but an
entire conidioma has not been drawn. In the absence of conidia, the conidio-
mata could scarcely, if at all, be distinguished from small conidiomata of K.
yuccifoliorum (Fig. 3, A). Conidiomata are 250-300 (-550) 4m diam and up to
450 ym tall. The wall is thick and cup-shaped. As described by Hohnel (4),
the top of a conidioma is composed of parallel filaments of elongate golden cells
which are very dark-walled in the area surrounding the ostiole. Microconidia
are formed on the upper conidiomatal wall and in the ostiolar channel. Conidia
are up to 80 ym long, considerably longer than the 50-61ym often cited (6, 13).
Part of this difference may be attributed to the irreversible shrinkage of conidia
from herbarium specimens (1) which were measured. The amount of
filamentous material making up the top of a conidioma is variable. Spermatia
are formed in 1-4 separate locules in the filamentous portion of a stroma, or in
the central locule with or without accompanying conidia; each locule has an
ostiole, or locules coalesce and release spermatia from a single ostiole.
Spermatiogenous cells are discrete to integrated on a 1-celled (usually)
conidiophore, cylindric to elongate-conical, phialidic, and 5-16 x 2.5-3.2 pm.
Spermatia are hyaline, bacillar, and 4-8 x 1.6 um.
Conidia germinated readily and formed colonies with white aerial, surface, and
subsurface mycelium. The reverse was pink. Within 6 weeks, conidiomata had
formed which contained typical Kellermania yuccigena conidia. Planistromella
uniseptate and K. yuccigena were therefore considered to be two phases of a
single holomorph.
DEVELOPMENT
Before the host epidermis is broken, a Kellermania conidioma consists of a cup-
shaped wall containing maturing macroconidia. The top of a conidioma is
made up of columns of elongate golden cells bounded internally by smaller,
hyaline cells. As conidiomata mature, the columns of elongate cells lengthen,
microconidium formation occurs in and near the ostiolar channel, and the cells
that will surround the ostiolar channel become very dark. The microconidio-
genous cells look much like periphyses in perithecial ascostromata, but form
microconidia as an added feature. The function of microconidia is not known.
A macroconidium develops as a simple outgrowth of a holoblastic conidiogenous
cell. In conidiomata potentially developing beyond macroconidium formation,
columns of cells are attached well down the lateral walls of the stromata (Fig. 3,
compare A, f, & D, d, t) reducing the size of the single, central locule. These
conidiomata may form spermatiogenous locules, ascogenous locules, or, very
rarely in Kellermania yuccigena, additional conidial locules in the columnar
filamentous portion. Spermatia are formed in the central locule with or without
macroconidia, in small spermatial locules that develop in the lateral upright
filaments, or at both sites. Frequently a conidioma forms spermatia in a single,
269
Ane
yh
Fig. 3. Kellermania yuccifoliorum. A. Small conidioma. B. Microconidia.
C. Spermatia and spermatiogenesis. D. Diagrams of stromata. d=columnar
filamentous portion of stroma developing into an area of spermatiogenesis,
e=host epidermis, f=columnar filamentous portion of stroma, o=dark-walled
cells around ostiole or developing ostiole, s=site of spermatiogenesis, t=portion
of stroma which may develop ascomata, u=site of conidiogenesis and
spermatiogenesis, w=wall. Standard line = A, 80 pm; B, C, 27 wm; D, 200 pm.
Fig. 4. A, B, Kellermania yuccifoliorum. A. Conidiogenesis and origin of
apical appendage. B. Conidia. C. Kellermania anomala conidia. D, E,
Kellermania yuccigena. D. Conidiogenesis and origin of apical appendage. E.
Conidia. Standard line = A, D, 27 pm; B, C, E, 40 um.
271
central locule, and then develops 1-4 sites in the columnar filamentous portion
where the cells become hyaline and subdivided. Spermatia adhere to the several
flexuous hyphae produced at each site.
Planistromella develops in pre-formed stromata which have already produced
conidia, spermatia, or both. Walls of the central locule are collapsed by
expansion of the ascogenous locules. The asci mature among cells of the copious
centrum tissues. These elongate cells, arranged in columns, may be laterally
compressed between maturing asci, but not much stretched. Dissolution of the
sterile cells begins at the upper end of the locule and proceeds downward. They
are still abundant as early-formed asci mature. In P. yuccifoliorum the
interthecial tissues are clearly filamentous as early-formed asci mature. In P.
uniseptata the interthecial tissues are irregularly filamentous, dissociating into
single cells or 2-3-celled series as the asci mature. In P. uniseptata the asci
elongate greatly, extending beyond the periphysate ostile for ascospore release.
Probably living mature P. yuccifoliorum asci elongate similarly, but this could
not be ascertained in the herbarium material which was mostly immature.
DISCUSSION
Beyond its similarity to Planistroma, Planistromella relationships are not clear.
(a) Planistroma asci are like those of the Botryosphaeriaceae (P. F. Cannon,
personal communication), as are the non-septate ascospores of Planistroma.
However, the type, Botryosphaeria dothidea (Mougeot ex Fries) Cesati & De
Notaris, has the Pleospora type of centrum development (7) which is not found
in Planistromella or Planistroma. (b) Several ascomycetes have 2-septate asco-
spores. It is improbable that the two ascospore septa are formed simultaneously
in Kiessleriella ocellata (Niessl) Bose or Paraphaeosphaeria michoti
(Westendorp) O. Eriksson which have brown ascospores, or Buergenerula Sydow
with hyaline ascospores, because of their relationships. Leptoguignardia Miller
has hyaline ascospores with two simultaneously-formed septa. None of these
fungi has a multilocular stroma, and the ascostromata are separate from the
conidiostromata so far as is known, though the two are found in the same
material for Leptoguignardia. (c) Planistromella is also distinct from the
known teleomorphs for Stagonospora (11) which belong to Leptosphaeria (7
species), Pleospora (2 species), and Didymella (1 species). These fungi have
unilocular pseudothecia, pseudoparaphyses that originate above the asci and
grow downward to the base of the locule, and, except for Didymella, colored
ascospores that are several septate or dictyosporous. According to existing
descriptions, ascospore septation is sequential, not simultaneous.
The Planistromella anamorph, Kellermania yuccifoliorum, would once have
been forced into Stagonospora gigantea. However, the two taxa can be
separated easily. Conidia of K. yucctfoliorum are nearly always 2-septate, and
commonly have a short apical appendage; Stagonospora gigantea conidia are
(3-)4-5(-7) septate, and lack an apical appendage. K. yuccifoliorum forms
272
microconidia near the ostiole and in the ostiolar channel; SS. gigantea has
periphysis-like cells present in these areas, but does not form microconidia. The
wall of Kellermania is gray-black when young, sometimes becoming dark brown
in age, whereas the wall of S. gigantea is brown.
The new anamorph taxon was clearly related to Kellermania and Piptarthron
by details of the life cycle and structure of the conidiomata. These genera are
distinguished by the presence of a single apical appendage on Kellermania
conidia. Conidial appendages vary in origin. To place the anamorph properly,
it was necessary to determine whether the small conidial appendages of the
taxon newly separated from S. gigantea resembled the larger conidial
appendages of existing Kellermanza species.
Kellermania yucctfoliorum conidia originate as simple outgrowths of the
holoblastic conidiogenous cells (Fig. 4, A). The cells elongate, often with little
increase in width, and in many of them the apex of the conidium becomes filled
with material that cannot be distinguished from the cell wall in unstained,
living cells. The conidia then increase in width, become septate, and enlarge
somewhat until they are mature. The occluded apex is the ‘apical appendage’
which retains the shape of the conidial initial at the time of appendage
formation. The apical cell may be truncate even when conidia are mature, and
the appendage becomes distinguishable from the cell wall. Origin of the apical
appendage is ‘cellular’ in the sense that it is not material extruded from a cell,
but it differs from the ‘cellular’ appendages of Heteropatella, Scolecosporiella,
and Chaetoconis which are apical cells drawn out into a filiform appendage.
Conidiogenesis was also examined in fresh collections of K. yuccigena (Fig. 4,
D) with newly-formed conidiomata. In unstained, untreated preparations,
developing conidia elongated to a sharp point bounded by a normal-appearing
cell wall. The lumen of the apex filled, and the cytoplasmic contents in the
narrowed apex were replaced. The wall of the pointed apex was separable from
the surrounding conidial sheath, but could not be distinguished from the
contents replacing the cytoplasm in the attentuated apex. No septum was ever
seen between the apex and the remainder of the conidium, nor did the.
cytoplasmic contents retract from the apex as in Monochaettellopsis (8). During
the filling of the apex, small stainable areas persisted for some time. The
appendages originated as cellular projections, as do those of conidia of
Monochaetiella hyparrheniae Castell (8), but the protoplasmic contents were
then replaced. The origin of K. yuccigena appendages seemed identical to,
though more elaborate than, the origin of K. yuccifoliorum appendages.
The above results are contrary to those of an earlier report (12) which describes
Kellermania yuccigena appendages as unstainable material in the apical region
of the most immature subulate conidia which becomes increasingly elongated as
the conidium matures. The fresh collections were put aside for later study to
try to reconcile the differences. When the collections were re-examined several
months later, not one sharply pointed conidium could be found with an open
273
narrowed apex. All apices were filled, making it appear that the appendage
originated outside the cell wall and increased in size as the conidium developed.
The reaction of the fungus to storage had obscured the true origin of the
conidial apical appendage. Since the conidial appendages of the new taxon
originated like those of K. yuccigena, it was designated a species of Kellermania,
and not considered to be a new, related genus.
The close relationship of Kellermania and Piptarthron was recognized by Hohnel
when he validated the genus Piptarthron (5). The teleomorphs for the two
genera are also much alike, but differ consistently in the species studied to this
point. Typical aseptate Planistroma ascospores, differing in size, have been
found in three of the four Piptarthron species whose teleomorphs have been
observed (9 and unpublished). Planistroma obtusilunatum Ramaley has asco-
spores that are shaped differently from the other Planistroma species (10), but
they are aseptate. The Planistromella teleomorphs for K. yuccigena, and K.
yucctfoliorum differ from Planistroma by their possessing septate ascospores.
The same is true for the teleomorph of K. anomala. The single observed ascus
of that species contained medianly 1-septate ascospores. A simplification would
be to include both anamorph genera in the earlier genus, Kellermania, and both
teleomorph genera in Planistroma. However, the generic limits are remarkably
consistent as they stand.
ACKNOWLEDGMENTS
I must express my appreciation to the curator of the Herbarium, University of
California, for making materials housed at that institution available to me. My
thanks also to Francis A. Uecker for the corrections and valuable suggestions
made in his review of the manuscript and his correcting of the Latin diagnoses,
and to Deborah Berrier for her advice, comments, and encouragement with the
statistics.
Literature Cited
1. Baral, H. O. 1992. Vital versus Herbarium Taxonomy: Morphological
Differences Between Living and Dead Cells of Ascomycetes, and Their
Taxonomic Implications. Mycotaxon 44: 333-390.
2. Farr, D. F., Bills, G. F., Chamuris, G. P., and Rossman, A. Y. 1989.
Fungi on Plants and Plant Products in the United States. The American
Phytopathological Society, 3340 Pilot Knob Road, St. Paul, Minnesota
55121, U.S.A.
3. Heald, F. D., and Wolf, F. A. 1911. New Species of Texas Fungi.
Mycologia 3: 522. We
4. Hohnel, F. v. 1915. Fragmente zur Mykologie no. 900. Uber die Gattung
Kellermania Ellis and Everhart. Sitzungsber. Kais. Akad. Wiss. Wien,
Math-Naturwiss. cl. Abt. 1, 124: 82-84.
274
jae
13.
Hohnel, F. v. 1918. Fungi imperfecti. Beitrage zur Kenntnis derselben.
Hedwigia 60: 129-209.
Morgan-Jones, G., Nag Raj, T. R., and Kendrick, B. 1972. Genera
Coelomycetarum. VI. Kellermania. Can. J. Bot. 50: 1641-1648.
Parguey-Leduc, A. 1966. Recherches sur L’Ontogénie et L’Anatomie
Comparée des Ascocarpes des Pyrénomycétes Ascoloculaires. Ann. Sci.
Naturelles, Bot., 12 ser. 7: 505-690.
Punithalingam, E. 1992. The Nuclei and Conidial Appendages in
Monochaetiellopsis species (Coelomycetes). Nova Hedwigia 54: 255-267.
Ramaley, A. W. 1991. Fungi of Yucca baccata. 1. Piptarthron
pluriloculare and Its Teleomorph, Planistroma yuccigena. Mycotaxon 42:
63-75.
Ramaley, A. W. 1992. Fungi From Yucca baccata 2. Planistroma
obtusilunatum sp. nov., and Its Anamorph, Piptarthron uniloculare sp.
nov. Mycotaxon 45: 449-460.
Sivanesan, A. 1984. The Bitunicate Ascomycetes and Their Anamorphs.
A. R. Gantner Verglag K. G., F19490 Vaduz.
Sutton, B. C. 1968. Kellermania and Its Generic Segregates. Can. J.
Bot. 46: 181-196.
Sutton, B. C. 1980. The Coelomycetes. Commonwealth Mycol. Inst.,
Kew, Surrey, England.
MY COTAXON
Volume XLVI, pp. 275-281 April-June 1993
4-O-METHYLLIVIDIC ACID, A NEW LICHEN DEPSIDONE
JOHN A. ELIX and DEBRA A. VENABLES
Department of Chemistry, The Faculties, Australian National University,
GPO Box 4, Canberra, ACT, 2601, Australia
ABSTRACT : The depsidone, 4-O-methyllividic acid
(8-hydroxy-3,4-dimethoxy-1-(2'-oxoheptyl)-6-pentyl-
11-oxo-11H-dibenzo|[b,e][1,4]dioxepin-7-carboxylic acid
(11) has been identified in the lichens Hypocenomyce
foveata, Hypotrachyna immaculata, H. livida and H.
osseoalba and was synthesized and characterized.
Introduction
Minor biosynthetic variations observed within a lichen genus or a
group of related species can produce a large number of structurally related
metabolites or a chemosyndrome (Culberson & Culberson 1976). The
structural variations within a chemosyndrome take several forms and in
the orcinol depsidones (1) these include :
i. variations in the length of the polyketide derived side chain R!.
ii. the degree of oxidation of this side chain (i.e. CH2COR' or CH2CH)R’).
iii. the degree of methylation of the phenolic groups (i.e. R?, R8 = Me or H).
iv. the presence/absence of an additional phenolic hydroxy group (or
derived methoxy group) in position 4.
Lichens of the genus Hypotrachyna, more particularly H. osseoalba
(Vain.) Park & Hale, H. livida (Taylor) Hale and H. immaculata (Kurok.)
Hale produce such a chemosyndrome, the following members of which
have been identified previously; norcolensoic acid (2), colensoic acid (3),
hydroxycolensoic acid (4), methoxycolensoic acid (5), physodic acid (6), 3-O-
methylphysodic acid (7) [unfortunately this compound was originally
termed 4-O-methylphysodic acid, using a now-obsolete numbering system]
oxyphysodic acid (8) and lividic acid (9) (Chester & Elix 1981; Culberson 1966;
Djura, Sargent et al. 1977a, 1977b; Elix & Engkaninan 1976; Elix & Johnston
1991).
Materials and Methods
The lichen fragments were freed as far as possible from obvious
organic substrate material and extracted with warm acetone for thin-layer
chromatography (TLC) or with warm methanol for high performance
276
O
Ry
O
ORg
O
RgO COoH
Rg Re
1
O
Ry
R30
Rg
Ry R3 R4 Re R7
2 Cs5Hi1 H H CsHi; 4H
3 CsHi1 Me H Csi ta abe
4 Cs5H11 Me OH Cs5H11 H
3) Cs5H11 Me OMe CsH11 H
6 CH2COCs5H;; H H Czlivin eee
| CH»COCsH;; Me H cH
8 CH»COCsH;; H OH Cs5H}] H
9 CH2COC5H;; Me OH Cs5H11 H
OM H
10 CH2COCS5H Me € CsH)1
le} CH2COC5H 1 H OMe Cs5H)1 CH»C.6Hs
12 #CHy»COCsH;; Me OMe Cs5Hj1 CH»2Ce6Hs
277
liquid chromatography (HPLC). Compounds were identified by TLC using
the methods standardized for lichen products (Culberson & Ammann, 1979;
Culberson & Johnson 1976, 1982; Elix, Johnston & Parker, 1987) and by
HPLC with retention index values (RI) calculated from benzoic acid and
solorinic acid controls (Feige, Lumbsch, Huneck & Elix, 1993). HPLC used a
Spectra System, a Perkin-Elmer HS-5C18 column and a spectrometric
detector operating at 254 nm with a flow rate of 1 ml/min. Two solvent
systems were used: 1% aqueous orthophosphoric acid and methanol in ratio
3:7 (A) and methanol (B). The run started with 100% A and was raised to
58% B within 15 min., then to 100% B in 30 min. and isocratic elution in
100% B for a further 10 min.
The following lichens were studied:
Hypocemomyce foveata Timdal
AUSTRALIA. Australian Capital Territory. On trunk of Eucalyptus
macrorhyncha in dry Eucalyptus woodland, Canberra Nature Park, Bruce
Ridge, 35°15'S, 149°05'E, J. A. Elix 27485, 261.1991 (ANUC). New South
Wales. At base of trunk of Eucalyptus macrorhyncha in dry Eucalyptus
woodland, Great Dividing Range, Tuena-Binda road, 30 km north-west of
Crookwell, 34°13'S, 149°20'E, J. A. Elix 27483 , 12.ix.1990 (ANUC).
Hypotrachyna immaculata (Kurok.) Hale
AUSTRALIA. Australian Capital Territory. On Leptospermum shrubs in
Eucalyptus woodland, trail to Booroomba Rocks, 11 km south-west of
Tharwa, 35°35'S, 149°02'E, J. A. Elix 4027, 16.ii.1977 (ANUC). REPUBLIC of
SOUTH AFRICA. Cape Province. On old oaks, Stormsrivier, Distr.
Humansdorp, O. Almborn 4133, 28.viii.1953 (ANUC).
Hypotrachyna osseoalba (Vain.) Hale & Park
AUSTRALIA. Queensland. On Alphitonia in disturbed rainforest,
Lannercost State Forest, Old Mill Road, 36 km WSW of Ingham, 18°45'S,
145°49'E, J. A. Elix 15609, 19.vi.1984 (ANUC). NEW ZEALAND. South
Island. On kahikatea in swampy forest, Little Ditch, 8 km north-east of
Waimangaroa, J. A. Elix 7273, 26.ii.1980 (ANUC).
The following authentic compounds have been synthesised or
isolated previously: norcolensoic acid (2) (Chester & Elix, 1981); colensoic
acid (3) (Djura & Sargent 1976); hydroxycolensoic acid (4) and
methoxycolensoic acid (5) (Djura, Sargent et al. 1977a,b); physodic acid (6), 3-
O-methylphysodic acid (7) and oxyphysodic acid (8) (Elix 1975) and lividic
acid (9) (Elix & Engkaninan 1976).
278
Synthesis of 4-O-Methyllividic acid (10)
Benzyl 4-O-Methyllividate (11)
Benzyl 4-methoxyphysodate (12) (Elix & Engkaninan 1976) (17.5 mg,
29.6 umole) was treated with an excess of ethereal diazomethane for a
period of 6 hours. The ethereal solution was treated with glacial acetic acid
(2 drops) and washed in turn with saturated sodium bicarbonate solution,
brine, and dried (MgSOq4). The residue obtained on evaporation of the
solvent was crystallized from light petroleum to yield benzyl 4-O-
methyllividate (11) (11.6 mg, 65%), as colourless needles m.p. 109.5-111°
(Found: mol. wt. 604.2670. 12C35!H49!6Oo, require mol. wt. 604.2672). 1H
n.m.r. (CDCl3) 6 0.78-0.91, m, CH2CH3; 1.21-1.62, m, CH2(CH2)3CH3; 2.55, t,
COCH2CHy9; 3.26, t) ArCH2CH 9; 3.76, s, 2H, ArCH2CO; 3.89, s, OCH3; 5.37, s,
COsCH); 6.56, 6.75; 2s, H2, H9; 7.41, m, C6Hs; 11.19, s; OH. Mass aeheeatss
m/z 605 (10%), 604 (M, 29), 513 (3), 469 (6).
4-O-Methyllividic Acid (10)
A solution of benzyl 4-O-methyllividate (11) (11.6 mg, 19. umole) in
ethyl acetate, containing 10% palladium-on-carbon (3.9 mg) was stirred in an
atmosphere of hydrogen at room temperature for 1.5 hours. The reaction
mixture was filtered to remove the catalyst and the catalyst washed with
ethyl acetate. The combined filtrate was then evaporated under reduced
pressure and the residue crystallized from dichloromethane/light
petroleum, to afford 4-O-methyllividic acid (10) (5.0 mg, 51%) as colourless
needles, m.p. 156.5-159° (Found: mol. wt. 470.2305. 12C271H34!6O7, requires
mol. wt. 470.2305). 'H n.m.r. (CDCl3) 6 0.88-0.94, m, CH2CH3; 1.24-1.65, m,
CH?2(CH2)3CH3; 2.57, t, COCH2CH?; 3.40, bt, ATCH2CH?; 3.92, s, ArCH2CO;
3:913.96, 25, OCH376.53,16776; 2s, H2, HI; 112-14, b, OL es Mass. spectrum
m/z 471 (32%), 470 (M, 100), 293 (16), 292 (42), 277 (38), 263 (16), 221 (12), 207
(14).
Results and Discussion
The lichen Hypotrachyna immaculata was shown by HPLC and TLC
to produce nine phenolic metabolites, eight of which corresponded to
known lichen substances. These included the common cortical depsides
atranorin and chloroatranorin, and the orcinol depsidones norcolensoic
acid (2), colensoic acid (3), physodic acid (6), 3-O-methylphysodic acid (7),
oxyphysodic acid (8) and lividic acid (9), all of which were readily identified
by comparison with authentic materials (Figure 1).
Subsequently we have undertaken the synthesis of 4-O-methyllividic
acid (10), and found that the TLC and HPLC behaviour of this compound to
be identical with that of the eighth phenolic substance present in H.
immaculata. This new depsidone has also been detected in several other
lichens which contain the colensoic acid - lividic acid chemosyndrome
ie
chemosyndrome including Hypocenomyce foveata Timdal and
Hypotrachyna osseoalba (Vain.) Park & Hale.
4-O-Methyllividic acid (10) was prepared by hydrogenolysis of
benzyl 4-O-methyllividate (11), which in turn was obtained by
methylation of benzyl 4-methoxyphysodate (12) (Elix & Engakinan 1976).
The standardized chromatographic data for these compounds are listed in
Table 1.
7 (ppb:
13
14
Figure 1. H.p.l.c. trace of Hypotrachyna immaculata (J. A. Elix 4027 in ANUC).
S = internal standard.
ACKNOWLEDGEMENTS
We thank the Australian Research Council for generous financial
support of this project; Mr. G. Deeble (HPLC), Ms. Caroline Barclay (HPLC)
and Ms. S. A. Elix (TLC) who assisted in the determination of the chemistry
of many specimens. A Commonwealth Postgraduate Award to (D.V.) is
gratefully acknowledged.
280
Table 1: Standardized Chromatographic Data for Hypotrachyna Metabolites
Standard Rg values (x 100) were determined in three independent t.l.c.
solvent systems: (A) toluene / dioxane / acetic acid (180 : 45 : 5); ( (B*)
hexane / t-butyl methyl ether / formic acid (140 : 72 : 18); (C) toluene /
acetic acid (170 : 30). HPLC retention index (RI) values (x 100) are relative
to solorinic acid and benzoic acid.
Compound Rr. A B* RI
Norcolensoic Acid (2) 28 27] 31 67.0
Colensoic Acid (3) 4] 66 47 85.0
Hydroxycolensoic Acid (4) 39 50 42 63.6
Methoxycolensoic Acid (5) 44 61 47 77.0
Physodic Acid (6) 25 35 18 63.3
3-O-Methylphysodic Acid (7) 39 45 42 73.3
Oxyphysodic Acid (8) 15 34 13 Sia
Lividic Acid (9) o2 37 31 56.8
4-O-Methyllividic Acid (10) 41 46 20 68.6
Atranorin (13) (Standard) 75 ris 79 76.3
Chloroatranorin (14) (Standard) 74 73 81 80.6
LITERATURE CITED
Chester, D. O. & Elix, J. A. (1981). New metabolites from Australian lichens.
Aust. ]. Chem. 34: 1507-1511.
Culberson, C. F. (1966). The structure of a new depsidone from the lichen
Parmelia livida. Phytochemistry 5: 815-818.
Culberson, C. F. (1972). Improved conditions and new data for the
identification of lichen products by a standardized thin-layer
chromatographic method. J. Chromatogr. 72: 113-125.
281
Culberson, C. F. & Ammann, K. (1979). Standardmethode zur Dtinnschicht-
chromatographie von Flechtensubstanzen. Herzogia 5 : 1-24.
Culberson, C. F. & Culberson, W. L. (1976). Chemosyndromic variation in
lichens. Syst. Bot. 1: 325-339.
Culberson, C. F. & Johnson, A. (1982). Substitution of methyl tert.-butyl
ether for diethyl ether in the standardized thin-layer chromatographic
method for lichen products. J. Chromatogr. 238: 483-487.
Djura, P. & Sargent, M. V. (1976). Depsidone Synthesis. VI. Colensoic acid.
Aust. J. Chem. 29: 1069-1077.
Djura,-Pisargent, aM V7 Bip) JAG Engkaninan, WU) Huneck, 5.) &
Culberson, C. F. (1977a). Depsidone synthesis. VII. Isolation and
structure determination of hydroxy- and methoxy-colensoic acids.
Synthesis of methyl methoxy-O-methylcolensoate. Aust. J. Chem. 30:
599-607!
Djura, P., Sargent, M. V., & Clark, P. D. (1977b). Depsidone synthesis. X.
Methoxy- and hydroxy-colensoic acids. Aust. J. Chem. 30: 1545-1551.
Elix, J. A. (1975). 2'-O-Methylphysodic acid and hydroxyphysodic acid: two
new depsidones from the lichen Hypogymnia billardieri. Aust. J.
Chem. 28: 849-858.
Elix, J. A. & Engkaninan, U. (1976). The structure of lividic acid, a
depsidone from the lichen Parmelia formosana. Aust. J. Chem. 29: 203-
207.
Elix, J. A. and Johnston, J. (1991). Additional lichen records for New
Zealand. 3. Further representatives in the family Parmeliaceae.
Australasian Lichenological Newsletter 28: 7-10.
Elix, J. A., Johnston, J. & Parker, J. L. (1987). A Catalogue of Standardized
Thin Layer Chromatographic Data and Biosynthetic Relationships for
Lichen Substances (Aust. Nat. University, Canberra).
Elix, J. A., Johnston, J. & Parker, J. L. (1988). A Computer Program for the
Rapid Identification of Lichen Substances. Mycotaxon 31: 89-99.
reice,. G. 5.) Lumbsch, H: 17, buneck\S> &jElix J. A.” (1993) The
identification of lichen substances by a standardized high-performance
liquid chromatographic method. J. Chromatogr. in press.
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MY COTAXON
Volume XLVII, pp. 283-284 April-June 1993
ANAEROMYCES, AN EARLIER NAME FOR RUMINOMYCES
Y. W. HO
Department of Biology, Universiti Pertanian Malaysia,
43400 Serdang, Selangor, Malaysia.
D. J. S. BARR
Centre for Land and Biological Resources Research,
Central Experimental Farm, Research Branch,
Agriculture Canada, Ont. Canada.
N. ABDULLAH and S. JALALUDIN
Department of Animal Science, Universiti Pertanian Malaysia,
43400 Serdang, Selangor Malaysia.
ABSTRACT
Ruminomyces Ho is placed into synonymy with
Anaeromyces Breton and comparison is made between
A. mucronatus and A. elegans (Ho) Ho comb. nov.
There were two taxa of anaerobic polycentric fungi described in
1990 from the rumen of cattle (Bos taurus). One was from France and
the other from Australia. The one from France was named
Anaeromyces mucronatus Breton while the other from Australia
was called Ruminomyces elegans Ho. In our opinion the two generic
diagnoses are essentially similar and the two genera are synonymous.
Thus, there is no justification for retaining them as two separate genera.
The generic name Anaeromyces has priority as it was published July
19, 1990 whereas Ruminomyces was published August 15, 1990.
However, the species are morphologically distinct and should be
retained as separate taxa. The following combination is therefore
proposed:
Anaeromyces elegans (Ho) Ho comb. nov.
= Ruminomyces elegans Ho (Ho etal. 1990).
Mycotaxon 38 : 398 - 404.
284
Although the morphology and size of the sporangia and
zoospores of A. elegans and A. mucronatus are somewhat similar, the
rhizomycelia of the two species are clearly different.
The rhizomycelium of A. elegans is composed of much-
branched, narrow to wide hyphae ranging from 0.5 - 10 um wide.
Many of the wide hyphae are sausage-shaped with constrictions at
regular or irregular intervals. Some hyphae also produce lobed or
bead-like structures, 1.5 - 14.5 um wide x 1.5 - 12.5 um high.
Frequently, fine pegs (about 0.5 um wide) are produced from the
centre of these lobed or bead-liked structures. The characteristics of
these structures appear to be similar to the penetrating structures
“appressoria" produced by anaerobic fungi colonising grass and straw
fragments in the rumen of cattle and swamp buffalo (Ho et al/., 1988a,
b; 1991). The hyphae of A. mucronatus, though with constrictions, do
not possess the lobed or bead-like structures. Although the difference
may eventually prove to be inconsequential, in our opinion, it is
preferable to retain the two species until the cultures are compared
under similar growing conditions and analysed by molecular analysis.
REFERENCES
Breton, A., Bernalier, A., Dusser, M., Fonty, G., Gaillard-Martinie, B. &
Guillot, J. (1990). Anaeromyces mucronatus nov. gen., nov.
sp. A new strictly anaerobic rumen fungus with polycentric
thallus. FEMS Microbiology Letters 70,177 - 182.
Ho, Y.W., Abdullah, N. & Jalaludin, S. (1988a) Penetrating structures
of anaerobic rumen fungi in cattle and swamp buffalo. Journal
of General Microbiology 134, 177 - 181.
Ho, Y. W., Abdullah, N. & Jalaludin, S. (1988b), Colonization of guinea
grass by anaerobic rumen fungi in swamp buffalo and cattle.
Animal Feed Science and Technology 22, 161 - 171.
Ho, Y. W., Bauchop, T., Abdullah, N. & Jalaludin, S. (1990).
Ruminomyces elegans gen. et sp. nov., a polycentric anaerobic
rumen fungus from cattle. Mycotaxon 38, 397 - 405.
Ho, Y. W., Abdullah, N. & Jalaludin, S. (1991). Fungal colonization
of rice straw and palm press fibre in the rumen of cattle and
buffalo. Animal Feed Science and Technology 34, 311 - 321.
MY COTAXON
Volume XLVII, pp. 285-293 April-June 1993
A NEW SPECIES OF PIROMYCES FROM
THE RUMEN OF DEER IN MALAYSIA
Y.W.HO
Department of Biology, Universiti Pertanian Malaysia,
43400 Serdang, Selangor, Malaysia.
D.J.S. BARR
Centre for Land and Biological Resources Research,
Central Experimental Farm, Research Branch,
Agriculture Canada, Ottawa, Ont. Canada.
N. ABDULLAH, S. JALALUDIN
Department of Animal Science, Universiti Pertanian Malaysia,
43400 Serdang, Selangor, Malaysia.
H. KUDO
Tropical Agriculture Research Centre, Tsukuba, Ibaraki,
Japan.
ABSTRACT
Piromyces minutus, a new species of anaerobic fungus from
the rumen of deer is described and illustrated. The fungus is
characterised by small sporangia, each with a wide apical
discharge pore, persistent sporangial wall and straight,
unbranched main rhizoid terminating in a sparsely-branched
rhizoidal system.
The name Piromonas communis Liebetanz was established in
1910 for a uniflagellated rumen organism presumed at that time to be
a flagellated protozoan. Later, Orpin (1977) discovered a rumen
fungal isolate with a vegetative phase which he associated with
Liebetanz’s Piromonas communis, but his isolate was clearly a
Chytridiomycete. Because it is not possible to know the true nature of
the organism Liebetanz observed, Gold et al. (1988) established a new
286
generic name, Piromyces, for Orpin’s fungus and the description was
later emended by Li et al. (1990). The name Piromonas could perhaps
have been validated and retained as it had been used in the literature
up to 1988. However, Piromyces is now generally accepted.
There are two other species of Piromonas that need mention.
These are Piromonas minima Liebetanz and Piromonas maxima
Liebetanz, but we regard these as ambiguous names. The descrip-
tions were based entirely on zoospore size and morphology (Liebetanz,
1910), and although it is possible that these were chytrids, it is net
possible to verify or relate them to any known rumen fungal species.
At present, four species have been formally described and
included in the genus Piromyces. They are P. communis Gold (Gold
et al., 1988, = Piromonas communis Sensu Orpin, 1977), P. mae Li (Li
et al., 1990), P- dumbonica Li (Li et al., 1990) and P. rhizinflata Breton
(Breton et al., 1991).
Recently, two morphological forms of Piromyces were isolated
over a two-month period from the rumen contents of Sika or Japanese
deer (Cervus nippon) fed guinea grass (Panicum maximum) in
Malaysia. One of the morphological forms was similar to P. communis
but the other had conspicuously smaller sporangia with persistent wall
following release of zoospores through a wide apical pore and is
thought to be a new species as it is distinctly different from the four
Piromyces species described.
The method for isolating the fungi from the rumen contents was
similar to that described by Ho & Bauchop (1991) except that glucose
agar instead of cellulose agar was used in the roll tubes. The media
for maintaining and culturing the isolates were glucose sloppy agar,
cellulose sloppy agar and liquid medium with Whatman No. 1 filter
paper or rice straw as carbon source (Ho & Bauchop, 1991).
Piromyces minutus Ho sp. nov. Figs. 1-14
Thallus monocentricus eucarpicus. Sporangium omnino endo-
genum, plerumque parvum denique ellipsoidale vel pyriforme vel
globosum, 8-25 um latum, 8.5-28 um longum, unico poro apicali
emiitenti lato et pariete sporangli persistenti, sed interdum grandius
denique globosum vel parum irregulare, 40-80 um diametro, 1-2 (uno
vel duobus) poris emittentibus. Axis rhizoidei singularis, aliquando 2-4
(duo vel quatuor). Rhizoideum principale sporangiorum parvorum
nonramosum, systeme rhizoidali parce ramoso terminans. Zoosporae
globosae 5.7-7.5 um diametro, uno interdum duobus raro quatuor
287
flagellis usque ad 31 um longis. Sporangia perdurantia ignota.
Species obligate anaerobia.
Thallus monocentric and eucarpic consisting of a sporangium
and filamentous rhizoidal system. Sporangia strictly endogenous,
predominantly small, ellipsoidal, pyriform or globose, 8 - 25 um wide x
8.5 - 28 um long with one wide, apical discharge pore and persistent
sporangial wall but occasionally larger, globose or slightly irregular, 40
- 80 um diameter with one or two discharge pores. Rhizoidal axis
single, occasionally two to four. Main rhizoid of small sporangia
usually unbranched, terminating in a sparely-branched rhizoidal
system. Zoospores globose, 5.5 - 7.5 um in diameter with one
flagellum, occasionally two, rarely four, up to 31 um long. Resting
spores unknown. Obligate anaerobe.
From rumen contents of Sika or Japanese deer (Cervus
nippon), Universiti Pertanian Malaysia, Serdang, Malaysia, 12 May,
1992, Y.W. Ho and D.J.S. Barr, D2 (UPM) holotypus.
The zoospores of P. minutus are globose (Fig. 1), 5.5- 7.5 um
in diameter when actively moving or stationary but occasionally
irregular and amoeboid just before encystment. They are usually
monoflagellated, occasionally biflagellated and rarely quadriflagellated.
The flagellum is up to 31 um long including a short whiplash. Shed
flagella frequently possess a bead-like structure (0.5-1.5 umin
diameter) at one end. Similar flagellar structure has been reported by
Barr et al. (1989) and Ho et al. (1993).
Germination of zoospores produces strictly endogenous
thallus development in all the four types of culture media used and in
cultures of different ages, including old cultures 4-5 days old. The
encysted zoospore expands uniformly and produces a germ tube
(Fig. 2) which develops into a main rhizoid. Two types of rhizoidal
systems are formed. In one type the main rhizoid is often straight and
unbranched for some length (up to 225 um) before branching to form a
sparse rhizoidal system (Fig. 3). It is narrow, 1.5 - 4.5 um in diameter,
usually quite even over its length but occasionally with swellings and
tightly constricted points (Figs. 4). The ends of the rhizoids are straight
with moderately blunt tips. Usually a single main rhizoid is produced
but occasionally two or more are formed (Figs. 5-6). This type of
rhizoidal system is predominantly formed in all the four culture media
used. In the other type of rhizoidal system, the main rhizoid is much
wider, 5.5 - 11.5 um in diameter, sometimes coiled and more densely
branched (Figs. 7-9). The ends of the rhizoids are usually straight
Fig. 1. Zoospore of Pminutus. Fig. 2. A germinating zoospore.
Fig. 3. Rhizoidal system showing straight and unbranched main
rhizoid terminating in sparsely-branched rhizoids. Diameter of the
main rhizoid is even without constrictions. Fig. 4. Main rhizoid with
swellings and constrictions. Figs. 5-6. Rhizoidal system with two and
three main rhizoids respectively. Bar = 20um.
289
and moderately blunt but occasionally they are rounded and slightly
expanded resembling small pegs (Fig. 10).
At maturity, the rhizoidal system is separated from the
sporangium by a septum at the neck (the point between the sporangium
and main rhizoid) or at the base of the sporangium.
The sporangium develops from the expansion of the encysted
zoospore. Sporangium shape is variable. Small sporangia are predo-
minantly formed. They are mostly ellipsoidal to pyriform, 8 - 25 um
wide x 8.5 - 28 um long, occasionally globose. They possess the
rhizoidal system with long, straight main rhizoid terminating with
sparse branches. Zoospores are discharged following dissolution of a
wide apical portion of the sporangial wall (Fig. 11). After discharge, a
persistent cup-shaped portion of the sporangial wall with a wide apical
pore or opening remains (Fig. 12).
In all cultures, there are invariably a small number of larger
sporangia which are globose, up to 80 um in diameter, or occasionally
slightly irregular. These larger sporangia frequently possess a denser
rhizoidal system and occasionally with a coiled main rhizoid. Zoospore
discharge is through one or two wide pores or openings (Fig. 13).
The sporangial wall is smooth (as seen after zoospore release),
two-layered and possibly thicker than in other species of Piromyces.
There is an outer layer that is continuous with the main rhizoidal wall
and an inner layer continuous with the septum. In sporangia following
discharge, the two layers are occasionally separated (Fig. 14), and the
septum can be seen as part of the inner layer. The persistence of the
wall following discharge can be considered a distinctive characteristic
of this species but the configuration of the layers, and continuity with
the septum, cannot be ascertained until more species of both aerobic
and anaerobic chytrids are studied with transmission electron micros-
copy. There are some differences in septum morphology between
different species of chytrids. Powell (1974) reported that in the aerobic
chytrid Entophylyctis variabilis Powell, a transverse septum forms by
centripetal accumulation of material on the walls of the rhizoids where
they join the sporangium (i.e. the port). In Triparticalcar arcticum Barr
there is a ring of material, different in composition to the wall, that
partially closes the port (Barr, 1970). In Spizellomyces spp. the neck
port is open (Barr, 1984). In contrast, Heath et al. (1983) reported that
in Neocallimastix frontalis a septum is laid down at sporogenesis, and
that the septum is continuous with the sporangial wall. A similar
continuity of septum and inner wall can be seen in the cellulolytic,
aerobic chytrid Rhizophlyctis rosea (de Bary & Woronin) Fischer
290
Figs. 7-9. P. minutus with wide main rhizoid and more densely-
branched rhizoidal system. The main rhizoid is sometimes coiled.
Note the ends of the rhizoids are straight and moderately blunt. Fig.
10. Rhizoidal system with densely-branched rhizoids and tips of lateral
rhizoids are rounded and slightly expanded. Bar = 20 um.
Fig. 11. Zoospores are released following the dissolution of a
wide apical portion of the sporangial wall. Fig. 12. A persistent cup-
shaped portion of the sporangium wall with a wide apical pore or
opening after discharge of zoospores. Fig. 13. An empty sporangium
with two wide pores or openings (arrows) through which zoospores are
discharged. Fig. 14. An empty sporangium after release of zoospores
showing a two-layed wall. The inner wall including the septum (arrows)
is detached from the outer wall which is continuous with the
main rhizoidal wall. Bar = 20um.
292
(unpublished). The septum in E. variabilis has plasmodesmata
whereas the septa in N. frontalis and R. rosea apparently do not.
Further studies of walls and septa in the Chytridiomycetes may reveal
homologies between the aerobic and anaerobic rumen taxa.
The presence of predominantly small sporangia each with a
wide apical pore or opening is the conspicuous feature of P. minutus
that distinguishes it from the other four species. In P. communis the
zoospores are released following overall dissolution of the sporangial
wall. For P. mae there are papillae that are thought to be products of
localised wall softening possibly similar to the more complex papillae in
other Spizellomycetales. Although rhizoid morphology is highly variable
within species of Chytridiomycetes, the straight, unbranched main
rhizoid terminating in a sparsely-branched rhizoidal system is very
characteristic of P minutus. This species also has strictly endogenous
sporangium which separates it from P. communis. All these charac-
teristics separate P. minutus from the other Piromyces species
described and thus require it to be assigned to a new species.
Other isolates examined: Five other isolates obtained from
the rumen contents of Timorensis deer (Cervus timorensis) at
Universiti Pertanian Malaysia, Serdang, Malaysia, November -
December, 1991, were also examined and their morphological
characteristics conform to the above description.
ACKNOWLEDGEMENTS
We thank Dr. J. Bisset for kindly providing the Latin translation
of the diagnosis, and Professor A. Nawawi for reviewing the manuscript.
We would also like to extend our thanks to Dr. M.K. Vidyadaran for
providing the Sika and Timorensis deer.
REFERENCES
Barr, D.J.S. (1970). Phlyctochytrium arcticum n. sp. (Chytridiales)
morphology and physiology. Can. J. Bot. 48, 2279-2283.
Barr, D.J.S. (1984). The classification of Spizellomyces, Gaertnerio-
myces, Triparticalcar and Kochiomyces (Spizellomycetales,
Chytridiomycetes). Can. J. Bot. 62, 1171-1201.
Barr, D.J.S., Kudo, H., Jakober, K.D. & Cheng, K.-J. (1989).
Morphology and development of rumen fungi: Neocallimastix
sp., Piromyces communis and Orpinomyces bovis gen. nov.,
sp. nov. Can. J. Bot. 67, 2815-2824.
is,
Breton, A., Dusser, M., Gaillard-Martinie, B., Guillot, J., Millet, L. &
Prensier, G. (1991). Piromyces rhizinflata nov. sp., a strictly
anaerobic fungus from faeces of the Saharian ass : a morpho-
logical, metabolic and ultrastructural study. FEMS Microbiol.
Lett. 82, 1-8.
Gold, J.J., Heath, |.B. & Bauchop, T. (1988). Ultrastructural description
of a new chytrid genus of caecum anaerobe, Caecomyces equi
gen. nov. sp. nov., assigned to the Neocallimasticaceae.
BioSystems 21, 403-415.
Heath, |.B., Bauchop, T. & Skipp, R.A. (1983). Assignment of the
rumen anaerobe Neocallimastix frontalis to the Spizellomyce-
tales (Chytridiomycetes) on the basis of its polyflagellate
zoospore ultrastructure. Can. J. Bot. 61, 295-307.
Ho, Y.W. & Bauchop, T. (1991). Morphology of three polycentric
rumen fungi and description of a procedure for the induction of
zoosporogenesis and release of zoospores in cultures. J. Gen.
Microbiol. 137, 213-217.
Ho, Y.W., Barr, D.J.S., Abdullah, N., Jalaludin, S. & Kudo, H. (19983).
Neocallimastix variabilis, a new species of anaerobic fungus
from the rumen of cattle. Mycotaxon 46, 241-258.
Li, J., Heath, |1.B. & Bauchop, T. (1990). Piromyces mae and
Piromyces dumbonica, two new species of uniflagellate
anaerobic chytridiomycete fungi from the hindgut of the horse
and elephant. Can. J. Bot. 68, 1021-1033.
Liebetanz, E. (1910). Die parasitischen protozoen der weiderkauer-
magens. Arch. Protistenkunde 19, 19-80.
Orpin, C.G. (1977). The rumen flagellate Piromonas communis: its
life-history and invasion of plant material in the rumen. J. Gen.
Microbiol. 99, 107-117.
Powell, M.J. (1974). Fine structure of plasmodesmata in a chytrid.
Mycologia 66, 606-614.
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Volume XLVI, pp. 295-357 April-June 1993
TYPE SPECIMENS OF LICHENS AND
LICHENICOLOUS FUNGI IN THE CANADIAN MUSEUM OF
NATURE (CANL)
PAK YAU WONG
Canadian Museum of Nature, P.O. Box 3443, Station 'D',
Ottawa, Ontario K1P 6P4, Canada
ABSTRACT
Type specimens of 737 lichen taxa and lichenicolous
fungi (82 holotypes, 418 isotypes, 1 lectotype, 3
lectoparatypes, 22 isolectotypes, 13 syntypes, 13 isosyntypes,
176 paratypes, 6 isoparatypes and 3 isoneotypes) in the lichen
herbarium of the Canadian Museum of Nature (CANL) are listed.
The name of the taxon, authority, literature citation,
category of type, locality information, collector, collector's
number, collecting date and CANL accession number are recorded
for each taxon, as well as, where applicable, exsiccat tittle
and number, and currently accepted name.
With the recent reorganization of the Canadian Museum of
Nature, a decision was made to upgrade the level of care for
the herbarium and to register all the type specimens in the
lichen collections (CANL).84Since thiss collection sis*ithe
largest in Canada and is one of the most important North
American lichen herbaria, the job was given the highest
priority.
The present catalogue lists in alphabetical order all
type specimens of lichens and lichenicolous fungi preserved
in the National Lichen Herbarium (CANL), which is housed at
the Canadian Museum of Nature. Authority abbreviations for
the epithets mainly follow Egan (1987) and its supplements
(Egan 1989, 1990). References for the original descriptions
are abbreviated according to Bridson (1991). The decision on
whether or not a particular specimen is actually a type was
made in the standard way, i.e., the label data were compared
with the protologue, and annotations of the original author
and subsequent annotators were considered. In some cases, the
type status was indicated in some way on the labels, but this
was not considered to be definitive. If the status of the
type is in question, e.g., because of a discrepancy in the
data, a query mark (?) is placed after the kind of type in
the catalogue. The label data, including locality
information, collector, collector's number and collecting
296
date are cited more or less as originally written on the
packet, with some standardisation, but the names of countries
are given in English reflecting current political boundaries
in so far as possible. The CANL accession number on the
packet is also presented. If the type is part of a published
exsiccata, the name of the series (with its author or
distributor) and number are also given at the end of the
entry. If paratypes occur in addition to other types or when
there is more than one paratype specimen, only locality,
collector, collector's number and CANL number are listed. No
attempt has been made to revise all the types by including
current synonyms, but where the type is known to be
synonymous with another name, the currently accepted name is
given.
I hope that this catalogue will be of assistance to
lichenologists who need to locate type specimens required for
their research. It may also help curators in other herbaria
locate types in their collections which may have gone
unnoticed.
This catalogue is not intended to be an authorative list
of the type status of each specimen, nor can it be a
definitive list of all the types now in CANL. As mentioned
above, further study may reveal that some specimens in the
list may not be types at all, and undoubtedly, some type
specimens exist in CANL that have been overlooked. Each
specialist must examine the evidence in order to decide the
status of the listed specimens and other CANL material not in
the list. Finally, it should be noted that in no case is any
new typification of any epithet being proposed in this
catalogue.
Acarospora canadensis H. Magn., Kongl. Svenska Vetensk. Acad.
Hand lo 73 232'"(1929)".
ISOTYPE. Canada: Ontario. Near the Hog's Back, Rideau River.
On limestone rocks. J. Macoun 3646, 2 May 1897 (CANL 11285).
Acarospora punae Lamb, Lilloa 14: 231 (1948).
ISOTYPE. Argentina: Prov. Tucuman, Valle de Tafi, western
Slope of Cumbre Potrerillo. On the overhung side of a large
schistose rock. I.M. Lamb 5290, 20 November 1947 (CANL
aa kee Bef) Be
Acolium bolanderi Tuck., Lich. of California Baier eereog).
ISOLECTOTYPE. United States: California, Oakland Hills. H.N.
Bolander 56, 1864 (CANL 68772). Reliq. Tuck. No. 101.
= Thelomma mammosum (Hepp in Hartung) Massal.
Alectoria ambigua Mot., Bryologist 67: 17 (1964).
ISOPARATYPES. Canada: Newfoundland, Trinity North District,
2.5 miles NNW of Northern Bight Sta. Exposed rock outcrop in
old fire-barren heath. T. Ahti 592, 15 August 1956. (CANL
52083); Placentia West, W of Swift Current, 2 miles NW of
Long Pond. On big boulder in treeless heath. T. Ahti S70 7.16
297
August 1956 (CANL 32344).
= Bryoria trichodes (Michx.) Brodo & D. Hawksw. subsp.
americana (Mot.) Brodo & D. Hawksw.
Alectoria arctica Elenk. & Savicz, Trudy Glavn. Bot. Sada 32:
730(1912)..
ISOTYPE. Russia: Novaya Zemlya, sinus Krestovaja. J.V.
Palibin, 1901 (CANL 104197).
= Evernia perfragilis Llano
Alectoria cornicularioides P. Jg@rg., Bryologist 78: 77
(1975).
ISOTYPE. China: Prov. Shensi, Kuan-tou-san. Among mosses. G.
Giraldi, 1896 (CANL 60512).
= Bryoria cornicularioides (P. Jgrq.) Brodo & D. Hawksw.
Alectoria corymbosa Hue, Expéd. Antarct. Franc. 9.0.3 29 O Ope bys
12 (1908).
ISOSYNTYPES. Antarctica: W. Graham Land (Palmer Peninsula)
Wandel (Booth) Island. Sur les roches. Expéd. Antarct. Fran¢.
272; 277 and 299, 25 November 1904 (CANL 5707, 5708).
= Catillaria corymbosa (Hue) Lamb
Alectoria fuscescens Gyelnik, Nytt Mag. Naturvidensk 70: 55
(1932).
ISOLECTOTYPES. Finland: Tavastia austr., Hollola, ad truncos
Pini locis apricioribus in silva. J.P. Norrlin, 18 September
1882 (CANL 29907 & 32345). Nyl. & Norrl.: Lich. Fenn. Exs.
No. 466.
= Bryoria fuscescens (Gyelnik) Brodo & D. Hawksw.
Alectoria glabra Mot., Fragm. Florist. Geobot. 6: 448 (1960).
ISOTYPE. United States: Washington, Clallum Co., Hurricane
Ridge, Olympic Peninsula. On trunk of Abies lasiocarpa. B.I.
Brown & W.C. Muenscher 129, 24 July 1950 (CANL 28849).
=Bryoria glabra (Mot.) Brodo & D. Hawksw.
Alectoria imshaugii Brodo & D. Hawksw., Opera Bot. 42: 59
(1977).
HOLOTYPE. United States: Idaho, Benewah County, ca. 4.5 miles
NW of St. Joe, top of St. Joe Baldy, near BLM Campground
Tingley Spring area. On Pseudotsuga menziesii in open
conifer stand. G.J. Schroeder L1944, 19 July 1971 (CANL
38827).
PARATYPES. United States: Calicornia, I.M. Brodo 20489 &
20502. (CANL 50581 & 50582). Oregon. L.H. Pike 1475 (CANL
42730).
Alectoria mexicana Brodo & D. Hawksw., Opera Bot. 42: 62
(1977).
HOLOTYPE. Mexico: Oaxaca, Sierra de San Felipe, in pino. On
pines. C.G. Pringle 195, 3 June 1894 (CANL 16582).
PARATYPES. Mexico: Oaxaca. J. Beharrel (CANL 55334, 55335 &
55342).
Alectoria stigmata Bystr., Fragm. Florist. Geobot. 20: 255
298
1974).
vaneeea United States: Alaska, Central Pacific Coast
District, Kenai Peninsula, Creek Ridge. H. Krog, 28 May 1957
(CANL 54995).
= Alectoria sarmentosa (Ach.) Ach. subsp. sarmentosa.
Amagdalaria consentiens var. japonica M. Inoue, J. Hattori
Bot. Lab. 56: 325 (1984).
ISOTYPE. Japan: Honshu, Pref. Nagano, Kiso-gun, Mitake-mura,
Mt. Ontake. On rock. M. Inoue 11474, 17 August 1976 (CANL
92651).
Amygdalaria continua Brodo & Hertel, Herzogia 7: 505 (1987).
HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands,
Moresby Island, Takakia Lake. At edge of water partially
submerged. I.M. Brodo 10880A, 4 July 1967 (CANL 70598)
PARATYPES. Moresby Island. I.M. Brodo 10880B, 10880D, 12804,
12819, 13960, 14288. (CANL 70599, 70601, 70605, 70553, 36494
& 70603).
Amygdalaria haidensis Brodo & Hertel, Herzogia 7: 508 (1987).
HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands,
Moresby Island, Tasu. On mined mountain behind Tasu camp,
forest line. On summit rocks. I.M. Brodo 12784, 28 July 1967
(CANL 70594).
PARATYPES. Moresby Island. I.M. Brodo 10834, 10933 & 14289.
(CANL 87857, 70604 & 70595).
Anaptychia fragilissima Kurok., Beih. Nova Hedwigia 6: 60
(1962).
ISOTYPE. Japan: Prov. Kii, Mt. Koya. On rocks. S. Kurokawa
60238, 17-18 December 1960 (CANL 20012). Kurokawa: Lich. Rar.
Crit EXs NO. 3.
Anaptychia japonica var. reagens Kurok., J. Jap. Bot. 35: 354
(1960).
ISOTYPE. Japan: Honshu, Prov. Sagami, Mt. Kintoki, Hakone. On
trunk of trees. S. Kurokawa 53064, 26 April 1958 (CANL
26391). Kurokawa: Lich. Rar. Crit. Exs. No. 53.
Anthracothecium japonicum Kashiw. & Kurok., J. Jap. Bot. 56:
308 (1981).
ISOTYPE. Japan: Hokkaido, Prov. Ishikari, Noborikawa, Yubari-
City. On bark of Acer. H. Shibuichi 6067, 4 August 1980 (CANL
78691). Kurokawa: Lich. Rar. Crit. Exs. No. 502.
Anzia afromontana R. Sant. in Moberg, Moberg: Lich. Sel. Exs.
Ups., Thunbergia 2: 1 (1986).
ISOTYPE. Tanzania: Arusha Prov., Mt. Meru, ca. 2 km N of
Kitoto Camp. On Stoebe kilimandscharica. R. Santesson 22949,
7 January 1971 (CANL 95077). Moberg: Lich. Sel. Exs. Ups. No.
Ls
Arthopyrenia submuriformis R. Harris, Michigan Bot. 12: 15
(1973).
ISOTYPE. United States: Pennsylvania, West Chester. On bark
Pe pe)
of hickory. F. Windle, July 1898 (CANL 19709). Cummings: Dec.
N. Amer. Lich. No. 299.
= Strigula submuriformis (R. Harris) R. Harris
Arthopyrenia willeyana R. Harris, Michigan Bot. 12: 16
(1973).
ISOTYPE. United States: Michigan, Iosco County. In jackpine-
aspen woods around Corsair State Forest Campground. R.C.
Harris 818, 18 September 1965 (CANL 37273).
= Anisomeridium nyssaiegenum (Ellis & Everh.) R. Harris.
Aspicilia quartzitica W. Weber, Bryologist 74: 183 (1971).
ISOTYPE. United States: Colorado, Boulder County, outer
foothills of the Front Range at Boulder. on resistant
quartzite rocks of the Lyons formation, base of the first
"Flatiron" on east face of Green Mountain. W.A. Weber, 24
January 1971 (CANL 52114). Weber: Lich. Exs. Colo. No. 364.
Bacidia colchica Vézda, Folia Geobot. Phytotax., Praha 14:
203 (1979).
ISOTYPE. Russia: Transcaucasia, Colchis. Distr. Gagra, in
valle angusta rivi Zo Ekvara. Ad folia Buxi colchicae. A.
Vézda, 13 June 1978 (CANL 73823). Vézda: Lich. Sel. Exs. No.
1660.
Bacidia lisowskii Vézda, Folia Geobot. Phytotax., Praha 15:
87 (1980).
ISOTYPE. Zaire: Haut-Zaire, Kisangani, prope vicum
Batiambale. Ad folia arborum. S. Lisowski, 27 October 1976
(CANL 73852). Vézda: Lich. Sel. Exs. No. 1690
Bacidia nivalis Follm., Philippia 4: 30 (1978).
ISOTYPE. United States: Washington, Whatcom Co., fleckférmig
an lichtoffenen, etwas windgeschiitzten, langfristig
schneebedeckten Kulm- und Neigungsflachen niederer Felsblicke
und bodennaber Gesteinstrtimmer im Sporastatietum testudineae
Frey, im Bereich der Sommerschneegrenze am Nordosthang des
Mount Baker liber dem Austinpass. G. Follmann, July 1969 (CANL
69925). Follmann: Lich. Exs. Sel. No. 262.
Bacidia scutellifera Vézda, Folia Geobot. Phytotax., Praha
10: 421 (1975).
ISOTYPE. Tanzania: Distr. Morogoro, montes Nguru, in monte
Mafulumula supra vicum Mnembule. Epiphylla in pluviisilva
montana. T. Pécs 6438, 20-21 August 1971 (CANL 57290). Vézda:
Lich. Sel. Exs. No. 1310.
Bacidia spirospora var. patagonica Lamb, Farlowia 4: 453
(1955).
HOLOTYPE. Argentina: Patagonia Prov. Chubut, Lago Cisnes near
Lago Menendez. On trunk of Colletia at edge of the forest.
I.M. Lamb 5933, 5.11.1950 (CANL 6024).
Bacidia vasakii Vézda, Folia Geobot. Phytotax., Praha 18: 64
(1983).
ISOTYPE. Russia: Colchis (Transcaucasia occid.), distr.
300
Adler, in valle angusta rivi Kudepsta, 5 km ab ostio in mare.
Ad folia Buxi colchicae. A. Vézda, 19 June 1979 (CANL 85982).
Vézda: Lich. Sel. Exs. No. 1869.
Bacidia violascens Kalb & Vézda, Folia Geobot. Phytotax.,
Praha 15: 309 (1980).
ISOTYPE. United States: Hawaii Insulae, Insula Kauai. Ad
ramulos arboris (Prosopis sp. cult.). O. Degener 34240, 29
February 1977 (CANL 76382). Vézda: Lich. Sel. Exs. No. 1733.
Baeomyces weberi Thomson, Bryologist 73: 632 (1970).
ISOTYPE. Papua New Guinea: Morobe District, Mount Kaindi near
Papua Wau. On clay banks of road-cuts not far below summit.
W.A. Weber, 18 June 1968 (CANL 33497). Weber: Lich. Exs.
Coloe No? 73.3137
Bahianora poeltii Kalb, Kalb: Lich. Neotr. Fasc. 8: 4 (1984).
ISOTYPE. Brazil: Bahia: Chapada Diamantina, Serra do
Tombador; etwa 1 km vor der Stadt Morro do ChapeG. Am Grunde
von Grdasern in einer Caatinga. K. Kalb, 20 July 1980 (CANL
89418). Kalb: Lich. Neotr. No. 302.
Bilimbia finkii Vainio, Mycologia 21: 35 (1929).
ISOTYPE. Puerto Rico: Rio Piedras. Open field on post. B.
Fink 534, 12 January 1915 (CANL 5847).
= Bacidia finkii (Vainio) Zahlbr
Bryoria carlottae Brodo & D. Hawksw., Opera Bot.: 42: 101
(1977).
HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands,
Graham Island, two mi. SE of Port Clements, S of the Tlell-
Port Clements Road. I.M. Brodo 18096, 11 July 1971 (CANL
38266).
PARATYPES. Canada: British Columbia, Queen Charlotte Islands.
I.M. Brodo 9900, 10657B, 12879, 18093, 18499 & 18503B (CANL
45536, 28902, 45490, 38264, 38265 & 42514).
Bryoria friabilis Brodo & D. Hawksw., Opera Bot.: 118 (1977).
ISOTYPE. United States: Washington, Gate. On oak trees. A.S.
Foster, 14 October 1911 (CANL 36151).
PARATYPES. British Columbia. I.M. Brodo 8179 (CANL 52607); J.
Macoun s.n. & 77 (CANL 16156 & 16162); Quebec. I.M. Brodo
18678 & 18691D (CANL 37596 & 37587).
Bryoria pikei Brodo & D. Hawksw., Opera Bot. 42: 125 (1977).
HOLOTYPE. United States: Oregon, Marion Co.. Epiphyte in an
old fruit orchard in a clearing in a douglas fir forest near
Winter Falls in Silver Creek Falls State Park E of Salem. On
young douglas fir. L. Pike 2475, 9 April 1972 (CANL 38271).
PARATYPES. Canada: British Columbia, Queen Charlotte Islands.
I.M. Brodo 12874A, 13839 & 18144A (CANL 45543, 45510 &
38383); Nova Scotia. J. Speer 73-24 (CANL 45159); United
States: Oregon. I.M. Brodo 20954 (CANL 99034); L.H. Pike 1110
(CANL 48810); Washington. I.M. Brodo 15438b (CANL 32327);
L.H. Pike 844 (CANL 48821.
301
Bryoria pseudocapillaris Brodo & D. Hawksw., Opera Bot. 42:
AA Sal @ EMA pe
HOLOTYPE. United States: Oregon, Curry County, Cape Blanco,
8 miles N of Port Orford. On Sitka spruce at headland. I.M.
Brodo 20539, 28 June 1974 (CANL 50596).
Bryoria salazinica Brodo & D. Hawksw., Opera Bot. 42: 130
(1977).
ISOTYPE. United States: Massachusetts, Springfield. W.G.
Farlow, May 1878 (CANL 36152).
PARATYPES. Canada: Prince Edward Island. J. Fabiszewski (CANL
31266); R.R. Ireland 10298 (CANL 35527).
Bryoria spiralifera Brodo & D. Hawksw., Opera Bot.: 42: 131
(1977).
HOLOTYPE. United States: California, Humboldt County. Pine
forest near Manila. On Pinus contorta. S. Dowty 137, 22
January 1972 (CANL 38403).
PARATYPES. United States: California. Brodo 20501, 20509 &
20511A (CANL 50580, 50594 & 50578).
Buellia excellens H. Magn., Medd. Gdteb. Bot. Trdadg. 17: 69
(1947).
ISOTYPE. Argentina: Prov. Salta, Quebrada de San Lorenzo. M.
Digilio-Grassi 335, 1946 (CANL 18205).
= Rhizocarpon compositum Lamb
Buellia galapagona W. Weber, Bryologist 74: 188 (1971).
ISOTYPE. Ecuador: Galapagos Islands, Isla Santa Cruz
(Indefatigable I.), Academy Bay. On lava rocks along the
shore of the bay just above high tide mark. W.A. Weber, 30
January 1964 (CANL 33508). Weber: Lich. Exs. Colo. No. 344.
Buellia imshaugii Hafellner, Beih. Nova Hedwigia 62: 58
(1979).
HOLOTYPE. Canada: South Saskatchewan. J. Macoun, 13 July 1879
(CANL 19326).
ISOTYPE. (CANL 19327).
Buellia multispora Kalb & Vézda, Folia Geobot. Phytotax.,
Praha 14: 203 (1979).
ISOTYPES. United States: Hawaii Insulae, Oahu, Keawaula
Valley. On living trunks of exotic Leucaena. O. & I. Degener
34257, 16 March 1977 (CANL 73863). Vézda: Lich. Sel. Exs. No.
1670; (CANL 102697). Kalb: Lich. Neotr. No. 408.
Buellia rivas-martinezii Barreno & Crespo, Philippia 2: 283
(1975).
ISOTYPE. Spain: Prov. Madrid, zerstreut an stark geneigten
kristallinen Gipsabbriichen in licht-und windoffenen
Trockenrasenlticken im Lecideion gypsicolae Crespo et Barr.,
620 m. SO, Cerros Yesiferos de Valdemoro nérdlich Aranjiez.
S. Rivas-Martinez & A. Crespo, March 1972 (CANL 57049).
Follmann: Lich. Exs. Sel. No. 143.
Buellia tephrodes Lamb, Farlowia 4: 468 (1955).
302
HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel
Huapi, Isla Victoria. On rocks at shore of lake, about 4 m
above water level. I.M. Lamb 5855 p.p. (CANL 18365).
Buelliella eximia Kalb & Hafellner, Kalb: Lich. Neotr. Fasc.
11: 6 (1990).
ISOTYPE. Venezuela: Lara Prov., Distr. Torres, Etwa 35 km E
von Barquisimeto. In einer trockenen Steppenlandschaft,
parasitisch auf Pyxine cocoes (Sw.) Nyl. K. & A. Kalb, 10
August 1989 (CANL 102740). Kalb: Lich. Neotr. No. 452.
Buelliella physciicola Poelt & Hafellner, Beih. Nova Hedwigia
623), 15S VOL979 )\.
ISOTYPE. Czechoslovakia: Moravia austro-occidentalis, in
valle fluminis Rokytna supra urbem MoravskYyY Krumlov. In
thallo Physciae sciastrae parasitice vigens, in rupibus
conglomeraticis. J. Poelt & A. Vézda, 16 April 1974 (CAN
73837). Vézda: Lich. Sel. Exs. No. 1675.
Byssoloma amazonicum Kalb & Vézda, Nova Hedwigia 51: 437
(1990).
ISOTYPES. Brazil: Amazonas, Regenwdlder am Rio Negro,
zwischen 100-200 km oberhalb von Manaus. Foliicola in
pluviisilva. K. Kaib, 14-18 October 1980 (CANL 103355).
Vézda: Lich Sel. Exs. No. 2427; (CANL 103753). Kalb: Lich.
Neotr. No. 482.
Byssoloma anomalum Kalb & Vézda, Nova Hedwigia 51: 438
(1990).
ISOTYPE. Brazil: Amazonas, Regenwdlder am Rio Negro, zwischen
100-200 km oberhalb von Manaus. K. Kalb, 14-18 October 1980
(CANL 103754). Kalb: Lich. Neotr. No. 483.
Calicium adspersum subsp. australe Tibell, Publ. Herb. Univ.
Uppsala 12: 1 (1984).
ISOTYPE. Australia: Tasmania, Ben Lomond National Park, 32 km
E of Evansdale, along Ben Lomond Road, 1 km NW of Carr Villa.
On decorticated, dead but still standing trunk of Eucalyptus
sp. L. Tibell 11465, 14 March 1981 (CANL 87449). Tibell:
Caliciales Exs. No. 76.
Calicium constrictum Tibell, Lichenologist 14: 223 (1982).
ISOTYPE. Costa Rica: Puntarenas, on old Camino Real, 3.5 km
SE of Paso Real. On bark of roadside fence-post. L. Tibell
8233, 30 December 1978 (CANL 87450). Tibell: Caliciales Exs.
NOW Lic
Calicium cryptocroceum Tibell, Publ. Herb. Univ. Uppsala 10:
L(1982)%
ISOTYPE. New Zealand: Canterbury, 9.5 km SSE of Arthur's
Pass, Bealy Spur, along track to Grasmere Hut. On
decorticated stump of Nothofagus solanderi v. cliffortioides.
L. Tibell 10084, 17 December 1980 (CANL 82033). Tibell:
Caliciales Exs. No. 51.
Calicium curtisii var. splendidula G. K. Merrill, Bryologist
303
LAS 1L07 C1909)
ISOTYPE. United States: Maine, Knox County, Rockland. On Rhus
typhina. G.K. Merrill, 3 May 1909 (CANL 20271). Merrill:
Tacn. Exse No. 724%
= Phaeocalicium curtisii (Tuck.) Tibell.
Calicium fuscipes Tuck., Genera Lichenum p. 240 (1872).
SYNTYPE. United States: New Jersey, Closter. C.A. Austin,
November 1864 (CANL 68717). Reliq. Tuck. No. 22.
= Mycocalicium fuscipes (Tuck.) Fink
Calicium obscurum G. K. Merrill, Bryologist 12: 107 (1909).
ISOTYPE. United States: Maine, Rockland. On dead fungus. G.K.
Merrill, 5 September 1909 (CANL 20337). Merrill: Lich. Exs.
NOs. 92%
= Chaenotheca brunneola (Ach.) Mull. Arg.
Calicium parvum Tibell, Symb. Bot. Upsal. 21: 84 (1975).
ISOTYPE. Canada: Ontario, Renfrew County, Petawawa Forest
Experiment Station, not far from Mill Lake. On trunk of Picea
in a swamp. L. Tibell 4638, 21 August 1972 (CANL 55417).
Caloplaca bisagnonis B. de Lesd., Bull. Soc. Bot. France 98:
he asf Ge toh i es
ISOTYPE. Italy: Genova, Val Bisagno, Prato. Sbarbaro, 21
April 1951 (CANL 17465).
Caloplaca brattiae W. Weber, Graphis Scripta 2: 168 (1989).
ISOTYPE. United States: California, Santa Barbara County,
Channel Islands, Santa Cruz Island. West end of the island.
On top of the rocky headland just above the spray zone. W.A.
Weber & C. Bratt, 8 January 1986 (CANL 97959). Weber: Lich.
Exs. Colo. No. 660.
Caloplaca britannica R. Sant., Lichenologist 24: 2 (1992).
ISOTYPE. Scotland: Caithness (V. C. 109), Crosskirk Bay, c.
10 km W of Thurso. On seashore rocks. R. Santesson 20410, 18
July 1969 (CANL 104124).
Caloplaca citrina var. arcis Poelt & Vézda, Vézda: Lich. Sel.
Exs. Fasc. 99: 6 (1990).
ISOTYPE. Austria: Stiria, Distr. Feldbach, Riegersburg. Ad
saxa andesitica sub arcem. G. Kantvilas, H. Mayrhofer & A.
Vézda, 22 September 1990 (CANL 103398). Vézda: Lich. Sel.
Exs. No. 2470.
Caloplaca fraxinea Lamb, Rep. (Annual) Natl. Mus. Canada,
Bill ewe ces 042"( 1954)",
HOLOTYPE. Canada: Nova Scotia, Cape Breton Island, at
Baddeck. On black ash bark in a swamp. J. Macoun 1737, 8 July
1898 (CANL 17744).
Caloplaca hensseniana Kalb, Kalb: Lich. Neotr. Fasc. 11: 7
(1990).
ISOTYPE. Venezuela: Lara Prov. Distr. Iribarreno, etwa 15 km
E von Barquisimeto. Epiphytisch in trockener Steppen-
304
Vegetation, untermischt mit Cacteen. K. & A. Kalb, 20 August
1989 (CANL 102742). Kalb: Lich. Neotr. No. 454.
Caloplaca litoricola Brodo, Bryologist 87: 98 (1984).
HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands,
Graham Island, Tow Hill. On shoreline rocks. I.M. Brodo 9905,
16 June 1967 (CANL 85435). Brodo: Lich. Canad. Exs. No. 156.
Caloplaca marina subsp. americana Arup, Bryologist 95: 158
(1992).
ISOTYPE. United States: Oregon, Lincoln County, Rocky Creek
Wayside State Park, just south of Whale Cove. On volcanic
seashore rocks. U. Arup 89189, 19 April 1989 (CANL 103721).
Caloplaca sbarbaronis B. de Lesd., Bull. Soc. Bot. France 96:
173 (1949).
ISOTYPE. Italy: Liguria orientalis, Bonassola. Rupicola prope
mare. Sbarbaro, February 1949 (CANL 17859).
Caloplaca sorediata var. tenuis H. Magn., Ark. Bot. 33A: 134
(1946).
SYNTYPE. Sweden: Lycksele Lappmark, par. Stensele,
Kyrkberget. Under overhanging rocks. A.H. Magnusson 7841, 5
July 1924 (CANL 17873).
= Xanthoria sorediata (Vainio) Poelt
Caloplaca spotornonis B. de Lesd., Bull. Soc. Bot. France
100: 177 (1953).
ISOTYPE. Italy: Savona, Spotorno, loco "Monte". Sbarbaro,
December 1951 (CANL: 17876).
Caloplaca trabicola H. Magn., Medd. Gdteb. Bot. Tradg. 18:
219 (1950).
PARATYPE. Uruguay: Florida Prov., Estancia, 25 de Agosto.
Sobre postes de alambrado. H.S. Osorio 1752, 22 November 1949
(CANL 17887).
Candelariella faginea Nimis & Poelt, Nova Hedwigia 49: 276
(1989).
ISOTYPE. Italy: Calabria, Serra del Prete, loco Massiccio del
Pollino dicto. Ad corticem Fagi. P.L. Nimis & J. Poelt, 10.
July 1988. (CANL 100651). Vézda: Lich. Sel. Exs. No. 2377.
Candelariella hudsonica Hakul., Ann. Bot. Soc. Zool.-Bot.
Fenn. "Vanamo" 27: 49 (1954).
HOLOTYPE. Canada: Northwest Territories, Franklin District.
On earth, north shore of Hudson Strait. R. Bell 1660, August
1897 (CANL 12961).
= Candelariella canadensis H. Magn.
Candelariella lambii Hakul., Ann. Bot. Soc. Zool.-Bot. Fenn.
"Vanamo" 27: 49 (1954).
HOLOTYPE. Argentina: Prov. Tucumen, Valle de Tafi, west slope
of Cumbre Potrerillo. On the top of a block of schistose rock
in alpine pasture. I.M. Lamb 5413, 24 November 1947 (CANL
12962).
305
Candelariella plumbea Poelt & Vézda, Folia Geobot. Phytotax.,
Praha 11: 88 (1976).
ISOPARATYPE. Austria: Karawanken, Karnten, Kanzianiberg bei
Finkenstein SE Villach, W- bis S-sei- tige Abbritiche. J.
Poelt, 10 February 1974 (CANL 73768). Poelt: Pl. Graecenses
No. 137.
Catapyrenium caeruleopulvinum Thomson, Bryologist 90:
30,1987.
ISOTYPE. United States: California, San Bernardino, 2 miles
SW of Parker Dam near the Colorado River. On soil. T.H. Nash
8449, 26 February 1974 (CANL 52409).
PARATYPE. United States: Arizona. T7.H. Nash 8047. (CANL
52394).
Catillaria italica B. de Lesd., Bull. Soc. Bot. France 97:
170 (1950).
ISOTYPE. Italy: Rapallo (Genova), + Loco opaco’_ infra
Montallegro, ad maceriam. Sbarbaro, November 1949 (CANL
5758).
Catillaria melanopotamica Lamb, Farlowia 4: 445 (1955).
HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel
Huapi, Isla Victoria. On trunk of Nothofagus dombeyi. I.M.
Lamb 5815, 26 January 1950 (CANL 5769).
PARATYPE. Argentina: Isla Victoria. I.M. Lamb 5836 (CANL
5768).
Cetraria alaskana cC. cCulb. & Culb., Bryologist 69: 200
(1966).
ISOTYPE. United States: Alaska, Along the Pitmegea River, 15
miles upstream from Cape Sabine. On wet tundra. J.W. Thomson,
10-17 July 1958 (CANL 20583). Thomson: Lich. Arct. No. 13.
= Cetrelia alaskana (C. Culb. & Culb.) Culb. & C. Culb.
Cetraria australiensis W. Weber ex Karnef., Bot. Not. 130:
yee ek: be
ISOTYPE. Australia: New South Wales, Snowy Mountains,
Kosciusko State Park. W.A. Weber & D. McVean, 2 February 1968
(CANL 52204). Weber: Lich. Exs. Colo. No. 454.
Cetraria fendleri f. coralligera W. Weber, Univ. Colorado
Stud., Ser. Biol. 10: 11 (1963).
ISOTYPE. United States: Colorado, Larimer County, Lumpy Ridge
nachst Twin Owls an der Route zum Gem Lake, 1.5 Meilen N Lake
Estes. R.A. Anderson, 27 August 1962 (CANL 71724). Crypt.
Exs. Vindob. No. 4439.
= C. coralligera (W. Weber) Hale
Cetraria iberica Crespo & Barreno, Anales Jard. Bot. Madrid
37: 205 (1980).
ISOTYPE. Spain: Guadalajara, Tamaj6én. Ad Cistus ladaniferus.
A. Crespo & E. Barreno, May 1977 (CANL 78851). Vézda: Lich.
Sel. Exs. No. 1752.
Cetraria islandica var. arborialis Zahlbr., Cat. Lich. Univ.
306
6: 333 (1930).
LECTOTYPE. Canada: British Columbia, Glacier, Marion Lake. On
bushes. J. Macoun, 23 August 1904 (CANL 15550).
= C. subalpina Imsh.
Cetraria subalpina Imsh., Mycologia 42: 746 (1950).
ISOTYPE. United States: Washington, Mt. Rainier National
Park, Mt. Wow. On branches of small shrubs. H.A. Imshaug
1876, 21 August 1948 (CANL 15549).
PARATYPES. Mt. Rainier National Park. H.A. Imshaug 261 & 267
(CANL 15551, 15552); Cataract Falls. H.A. Imshaug 510 (CANL
24532).
Chaenothecopsis nana Tibell, Publ. Herb. Univ. Uppsala 4: 4
(1979).
PARATYPE. Sweden: Varmland, Norra Finnskoga par., N of
Digerfallet. On trunk of Picea abies. S. Sundell 13382, 16
August 1979 (CANL 71126). Tibell: Caliciales Exs. No. 35.
Chaenothecopsis nigropedata Tibell, Symb. Bot. Upsal. 27: 135
(1987).
ISOTYPE. New Zealand: South Island, Southland, Longwood State
Forest, 19 km NNW of Riverton, along Pourakino River, N of
Pourakino Campground. On decorticated stump. L. Tibell 10428,
16 January 1981 (CANL 99793). Tibell: Caliciales Exs. No.
160.
Chaenothecopsis sagenidii Tibell, Symb. Bot. Upsal. 27: 148
(1987).
ISOTYPE. New Zealand: Otago, 15 km WNW of Owaka, Catlins
Forest, along Catlins River, S of the junction with Chloris
stream. On still corticated parts of decaying stump of
Nothofagus menziesii. L. Tibell 10282, 11 January 1981 (CANL
99795). Tibell: Caliciales Exs. No. 162.
Chaenothecopsis tasmanica Tibell, Publ. Herb. Univ. Uppsala
16: 6 (1985).
ISOTYPE. Australia: Tasmania, Mt. Field National Park, W
shore of Lake Dobson. On dead leaves of Richea pandanifolia.
Lb. .Tibeli 11227, (¢S March’ ©1981) '*-(CANL” °89742))."" *Tibely:
Caliciales Exs. No. 118.
Charcotia rufidula Hue, Bull. Soc. Bot. France 62: 17 (1915).
ISOTYPE. Antarctica: Graham Land (Palmer Peninsula), Booth
(Wandel) Island. Deux Exped. Antarct. France 123, October
1908. (CANL 10604).
= Arthonia rufidula (Hue) D. Hawksw., R. Sant. & @vstedal
(1991).
Cheiromycina petri D. Hawksw & Poelt, Lichenologist 22: 219
(1990).
ISOTYPE. Austria: Styria, in collibus "Oststeierisches
Hiigelland", distr. Feldbach, Lamberg, in pago Petersdorf II.
Ad corticem trunci Betulae. J. Poelt & H. Pittoni, 27
November 1987 (CANL 103356), Vézda: Lich. Sel. Exs. No. 2428.
307
Cladina conspicua Ahti, Ann. Bot. Fenn. 23: 224 (1986).
ISOTYPE. Canada: Newfoundland, Humber East District, ca. 2.4
km N of Gaff Topsail, timberline heath. 7. Ahti 2997, 27 June
1956 (CANL 96933).
PARATYPE. Canada: Newfoundland, St. Mary District. T. Ahti
9461. (CANL 96934).
Cladonia alpestris f. aberrans des Abb., Bull. Soc. Sci.
Bretagne 16, Fasc. hors sér. 2: 93 (1939).
ISOLECTOTYPE. United States: Alaska, St. Paul Island. J.
Macoun, 26 June 1897, (CANL 6540). Macoun: Canad. Lich. No.
LVL;
Cladonia anitae Culb. & C. Culb., Mycologia 74: 663 (1982).
ISOTYPE. United States: North Carolina, Onslow Co., inter
Folkstone et Holly Ridge. Ad terram arenariam. W.L. Culberson
18539, 1 April 1981 (CANL 85967). Vézda: Lich. Sel. Exs. No.
1854.
Cladonia arbuscula subsp. beringiana Ahti, Ann. Bot. Soc.
Z001.-Bot. Fenn. "Vanamo" 32: 109 (1961).
ISOTYPE. Canada: Northwest Territories, Mackenzie District,
Great Bear Lake, at head of Great Bear River where it flows
out of Great Bear Lake. A.A. Lindsey 417, 23 July 1951 (CANL
9766).
= Cladina arbuscula (Wallr.) Hale & Culb. subsp. beringiana
(Ahti) Golubk.
-Cladonia boliviana Ahti, Ann. Bot. Soc. Zool.-Bot. Fenn.
wVanamno™ (3227 °PS i (1L96L)s
ISOTYPE. Bolivia: La Paz, 10,000 ft. M. Bang 20a (20 on
holotype), 1889. (CANL 59522).
= Cladina boliviana (Ahti) Ahti
Cladonia convoluta var. vagans Follm., Philippia 2: 208
(1975).
ISOTYPE. Spain: Prov. Teruel. Zwergstrauchheide in der Sierra
de la Costera unweit Fuentes Calientes. G. Follmann, July
1973 (CANL 57051). Follmann: Lich. Exs. Sel. No 145.
Cladonia crispata var. infundibulifera f. albopunctata G. kK.
Merrill, Bryologist 11: 110 (1908).
SYNTYPES. Canada: Yukon, Hunker Creek. In bogs. J. Macoun 104
& 108, 28 July 1902 (CANL 7411 & 7382).
= C. crispata (Ach.) Flotow var. cetrariiformis (Delise)
Vainio (fide T. Ahti)
Cladonia favillicola Trass, Investigationes Naturae Orientis
Extremis (Tallinn) p. 198 (1963).
ISOTYPE. Russia: Kamczatka, 20 km a Kozyrevsk in SO, ad
flumen Pahtza, in lariceto-ledose, ad laricis truncum
prolapsum crasso strato ex favilla vulcanica et aculeis
laricis tectum. H. Trass, 23 August 1960 (CANL 57970).
= C. botrytes (K. Hagen) Willd.
Cladonia gracilis subsp. vulnerata Ahti, Ann. Bot. Fenn. 17:
308
207 (1980).
ISOTYPE. United States: Alaska, Chugach Mts., Blueberry Lake
Campground, Mile 23, Richardson Hwy. In montane tundra. T.
Ahti & J.W. Thomson 23642, 25-26 July 1967 (CANL 75105).
PARATYPES. Canada: British Columbia. I.M. Brodo 9760 & 18156.
(CANL 31098 & 61622).
Cladonia hedbergii Ahti, Lichenologist 9: 4 (1977).
ISOTYPE. Kenya: Central Prov., Nanyuki Distr., Mt. Kenya,
National Park Road (Naro Moru Track). On the ground in a wet
slope in the Hagenia-Hypericum zone. R. Santesson "32495"
(22040 in protologue), 23 January 1970 (CANL 99586). Moberg:
Lich. Sel. Exs. Upsal. No. 56.
Cladonia kauaiensis G. K. Merrill, Ark. Bot. 31A(6): 28
(1944).
ISOLECTOTYPE. United States: Hawaiian Islands, Kauai. On
earth. A.A. Heller, 10-16 September 1895 (CANL 20334).
Merrill: Lich. Exs. No. 89.
Cladonia labradorica Ahti & Brodo, Bryologist 84: 238 (1981).
HOLOTYPE. Canada: Québec, Post-de-la-Baleine, 8 milles de la
mer (Baie d'Hudson). A. Vachon, 20 August 1974 (CANL 71500).
PARATYPES. Canada: Quebec. J.P. Ducruc 73-4062 (CANL 49135);
G. Lemieux 21399 (CANL 70052); J.L. Lethiecgq QFB-E 4495 &
4497 (CANL 48848 & 48881).
Cladonia libifera Savicz, Novosti Sist. NizSih Rast. 1965:
167 (1965).
ISOTYPE. Russia: Sibiria orientalis, Respublica autonoma
Jakutensis, in valle fluminis Aldan, prope Chandyga apud
fontem "Tjoply kljucz' in lariceto ad truncum putridum. L.N.
Tjulina, 1949 (CANL 48266).
= C. pocillum (Ach.) O. Rich.? (fide T. Ahti)
Cladonia magyarica Vainio, Fl. Hung. Exs. No. 715 (1927).
ISOLECTOTYPE. Hungary: Comit, Pest Prov., "Bugaci nagyerddé"
prope oppidum Kecskemét. Ad arenam mobilem in silva. G.
TimkO, May 1924 (CANL 19843).
Cladonia norvegica Tonsberg & Holien, Nordic J. Bot. 4: 79
(1984).
ISOTYPE. Norway: S@r-Tr@ndelag, Melhus, inter montem Loasen
et rivulum Loa. Ad basin abscisam arboris (Picea abies). T.
Teansberg 6870, 5 June 1982 (CANL 89184). Vézda: Lich. Sel.
Exs. No. 1978.
Cladonia patagonica A. Evans, Rev. Bryol. Lichénol. 24: 135
(1955).
ISOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Menéndez,
W. end. I.M. Lamb 5886, 2 February 1950 (CANL 9081).
= C. squamosa (Scop.) Hoffm. (barbatic acid chemotype) (fide
TRAnt sw 992)
Cladonia physodalica Elix, Lich. Austral. Exs. Fasc. 9, No.
204 (1990).
309
ISOTYPE. Australia: Queensland, Main Coast Range, Mt. Lewis
Track, Mary Creek, 15 km NNW of Mt. Molloy. On soil on large
boulder. J.A. Elix 16947, 30 June 1984 (CANL 102793). Elix:
Lich. Austral. Exs. No. 204.
= C. poeciloclada des Abb. (physodalic acid chemotype) (fide
T. Ahti 1992).
Cladonia rangiferina f. caerulescens Schade, Ber. Deutsch.
Bot. Ges. 69: 284 (1956).
ISOTYPE. Canada: Manitoba, Oxford Lake. H.J. Scoggan 60, 3
July 1949. (CANL 9394).
= Cladina stygia (Fr.) Ahti
Cladonia rangiferina f. leucosticta G. K. Merrll, Bryologist
11: 109 (1908).
SYNTYPES. Canada: Yukon, Hunter Creek. In a bog. J. Macoun
112 & 113 (CANL 9141 & 9140).
= C. wainii Savicz (fide T. Ahti)
Cladonia subchordalis A. Evans, Rev. Bryol. Lichénol. 24: 133
(1955).
ISOTYPE. Argentina: Patagonia, Prov. Chubut, w. end of Lago
Menéndez, Torrecillas Stream. On mossy soil between morainic
boulders. I.M. Lamb 5917, 4 February 1950 (CANL 9671).
Cladonia terrae-novae Ahti, Arch. Soc. Zool.-Bot. Fenn.
"Vanamo" 14: 131 (1960).
ISOTYPE. Canada: Newfoundland: Placentia West District, 4
miles NW of Long Pond. T. Ahti 108, 16 August 1956 (CANL
58385).
= Cladina terrae-novae (Ahti) Hale & Culb.
Cladonia terrae-novae f. cinerascens Ahti, Ann. Bot. Soc.
Zool.-Bot. Fenn. "Vanamo" 32: 82 (1961).
ISOTYPE. Canada: Newfoundland, Placentia West District, 2 mi.
NW of Long Pond. T. Ahti 7, 17 August 1956 (CANL 58408).
= Cladina terrae-novae (Ahti) Hale & Culb. f. cinerascens
(Ahti) Brodo
Cladonia thomsonii Ahti, Bryologist 81: 334 (1978).
ISOPARATYPE. United States: Northern Alaska, on the point of
Point Barrow. J.W. Thomson, S. Shushan & A.J. Sharp, 18 July
1958 (CANL 20591). Thomson: Lich. Arct. No. 21.
Cliostomum luteolum Gowan, Mycologia 82: 769 (1990).
HOLOTYPE. Canada: New Brunswick, Albert Co., Fundy National
Park, Point Wolfe. On Picea rubens. S.P. Gowan 3836, 1 August
1980 (CANL 82486).
Cliostomum vitellinum Gowan, Mycologia 82: 769 (1990).
PARATYPE. Canada: New Brunswick, Westmoreland Co., near
Sackville, Johnson's Mills. On trunk of Abies balsamea. M.
Fredericksen 78008, 1 February 1978 (CANL 67236).
Coccocarpia stellata Tuck., Proc. Amer. Acad. Arts. 5: 402
(1862).
310
ISOTYPE. United States: South Carolina, Santee Canal. H.W.
Ravenel 250 (CANL 68742). Reliq. Tuck. No. 63.
Coccotrema maritimum Brodo, Bryologist 76: 263 (1973).
HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands,
Moresby Island, Jedway. On shoreline rocks. I.M. Brodo 12459,
25 July 1967 (CANL 30581).
PARATYPES. Canada: British Columbia. I.M. Brodo 14355 (CANL
30862); K.E. Ohisson 1011, 1023) 1050, ) 1059, 21/3 i216
2448; (CANL 32299, 32300, 32295) 32296732298 ,32297) &°32301)7
G.F. Otto 1504 (CANL 36507); M.J. Shchepanek 253 (CANL
30876).
Collema microptychium Tuck., Lich. Calif.: 35 (1866).
ISOLECTOTYPE. United States: Massachusetts, Amherst,
Parttrigg's Swamp. E. Tuckerman, 1861 (CANL 23165).
= C. leptaleum Tuck.
Coniocybe amabilis Tibell, Publ. Herb. Univ. Uppsala 10: 6
(1982).
ISOTYPE. New Zealand: Wellington, Tongariro National Park, 3
km NE of Ohakune, along Mountain Road. On decorticated stump
of Dacrydium cupressinum. L. Tibell 13605, 3 June 1981 (CANL
82046). Tibell: Caliciales Exs. No. 65.
Conotremopsis weberiana Vézda, Folia Geobot. Phytotax., Praha
1281411997) 2
ISOTYPE. Australia: Tasmania, Lake St. Clair National Park.
On saplings near shore of Lake St. Clair at beginning of
trail to Mt. Hugel. W.A. Weber & D. McVean 49678, 24 February
1968 (CANL 92385).
Dactylina beringica Bird & Thomson, Canad. J. Bot. 56: 1612
(1978).
ISOTYPE. Canada: British Columbia, Summit Lake. Mile 392.5,
Alaska Hwy, slope of Mt. George. J.W. Thomson & T. Ahti
16553, 13 August 1967 (CANL 59778).
Dermatocarpon corticola ("corticolum") Rd&dsdnen, Anales Soc.
Ci. Argent. 128: 147 (1939).
ISOTYPE. Uruguay: Dept. Artigas, Sucio. On bark of trees.
W.G. Herter, 1934 (CANL 1063).
Dermatocarpon gorzegnoense Servit, Ann. Mus. Civico Storia
Nat. Giacomo Doria 66: 242 (1953).
ISOTYPE. Italy: Piemonte, Langhe, Gorzegno. Sbarbaro,
September 1951 (CANL 1081).
Dermatocarpon vagans Imsh., Mycologia 42: 753 (1950).
ISOTYPE. United States: Wyoming, NE corner of Yellowstone
National Park. On arid soil. H.A. Imshaug 15, 23 June 1948
(CANL 1185).
Dimerella chiodectonoides Kalb, Kalb: Lich. Neotr. Fasc. 10:
7 (1988).
ISOTYPE. Ecuador: Azuay Prov., etwa 35 km siidlich von Cuenca.
311
In sehr dichten, feuchten Waldresten in einem Graspdramo. K.
& A. Kalb, 26 August 1987 (CANL 102706). Kalb: Lich. Neotr.
No. 417.
Dimerella degeneri Kalb & Vézda, Folia Geobot. Phytotax.,
Prana 115253107 (1980).
ISOTYPE. United States: Hawaii Insulae, Insula Oahu,
Kaukonahua Gulch prope territorium Universitatis. Ad corticem
arboris (Ficus sp. cult.). O. et I. Degener 34269, 17 May
1977 (CANL 76378). Vézda: Lich. Sel. Exs. No. 1729.
Dimerella frederici Kalb, Folia Geobot. Phytotax., Praha 15:
310 (1980).
ISOTYPES. United States: Hawaii, island Hawaii, Kilauea, Kau
Desert. On bark of naturalized Samanea saman. O. et I.
Degener 34230, 1 February 1977 (CANL 76379, 77581, 77633, and
VODs \ovanveZda isi Chace se ExXGs NOt 6173074 Poelt:) PL.
Graecenses No. 200; Follmann: Lich. Exs. Sel. No. 343; Kalb:
hich.’ Neotr.s)No. 419,
Dimerella isidiigera Vézda & Osorio, Vézda: Lich. Sel. Exs.
Masco 4 2762 7(1989))..
ISOTYPE. Uruguay: Montevideo, Parque Rivera. Ad corticem
Cupressi loco umbroso. H.S. Osorio, 12 October 1988 (CANL
100603). Vézda: Lich. Sel. Exs. No. 2329.
Dimerella pocsii Vézda & Farkas, Folia Geobot. Phytotax.,
Prahay232) 193.7(1988)*
ISOTYPE. Tanzania: Tanga regio, Usambara Orientalis,
reservatum naturae Kihuhwi Forest dictum, secus trametem
inter Kihuhwi et Wamkoro. Foliicola in pluviisilva. E. Farkas
86228, 3 November 1986 (CANL 97332). Vézda: Lich. Sel. Exs.
Nowe2230.
Dimerella subdentata Vézda & Thor, Vézda: Lich. Sel. Exs.
Fasc. 94: 3 (1989).
ISOTYPE. Australia: Queensland, circa 15 km ad occidentem et
septentriones a Tully prope litorem Mission, secus Stony
Creek. Ad corticem arboris in pluviisilva. G. Thor 5721, 28
November 1985 (CANL 100607). Vézda: Lich. Sel. Exs. No. 2333.
Dimerella subfallaciosa Vézda & Farkas, Vézda: Lich. Sel.
Exo. erasc .0:98's)2-7(01990).
ISOTYPE. Tanzania: Regio Morogoro, montes Nguru, in valle
"Dikurura", 3 km ad orientem a "Mhonda Mission". In
pluviisilva montana. E. Farkas 89100, 22 March 1989 (CANL
103357). Vézda: Lich. Sel. Exs. No. 2429.
Dimerella usambarensis Vézda & Farkas, Folia Geobot.
Pauytotax., Praha 23, 196 9(1988)..
ISOTYPE. Tanzania: Tango regio, Usambara Orientalis,
reservatum naturae Marvera Forest dictun, 6 km ad
septentriones et orientem ab Amani. Foliicola. E. Farkas & T.
Pécs 86228, 12 December (November in protologue) 1986 (CANL
67333) .)VézdasiLichwesel J iExe si Na. i2230".
Die
Diploschistes montevidensis H. Magn., Medd. Gdteb. Bot.
Trddg. 18: 223 (1950).
ISOTYPE. Uruguay: Montevideo, Pajas Blancas. Sobre la tierra.
H.S. Osorio 1621, 20 February 1949 (CANL 2236).
= D. conceptionis Vainio
Diploschistes sbarbaronis B. de Lesd., Bull. Soc. Bot. France
99: 147 (1952).
ISOTYPE. Italy: Toscana, Siena, Latericola in villa Solaia
(Monteliscai). Sbarbaro, May 1951 (CANL 2241).
= D. actinostomus (Pers. ex Ach.) Zahlbr.
Diploschistes straminescens Zahlbr., Mycologia 22: 71 (1930).
ISOLECTOTYPE. Puerto Rico: Mayaguez. Open road-side on clay
bank. B. Fink 1016, 18 December 1915 (CANL 2279).
= D. hypoleucus Zahlbr.
Dirina californica Tuck., Lich. Calif. p. 422 (1866).
ISOLECTOTYPE. United States: California, Oakland. H.N.
Bolander 129, 1866 (CANL 68755).
= Schismatomma californicum (Tuck.) Zahlbr.
Dolichocarpus chilensis R. Sant., Svensk Bot. Tidskr. 43: 552
(1949).
ISOTYPE. Chile: Prov. Coquimbo, Coquimbo. On spines of Cereus
in dry hill near the sea-shore. R. Santesson 2620a, 1 August
1940 (CANL 2113).
Endocarpon mailae Lamb, Lilloa 14: 213 (1948).
PARATYPE. Argentina: Prov. Tucuman, Sierra de Javiev, E.
side. On rocks at roadside. I.M. Lamb 5096, 14 September 1947
(CANL 1197).
Endocarpon tuckermanii Rav. ex Mont., Sylloge Gener. Spec.
Cryptog.: 359 (1856).
ISOTYPE. United States: South Carolina, Santee Canal. On
Carya sp. among mosses. H.W. Ravenel 138 (CANL 1180). Reliq.
Tuck. No. 42.
= Catapyrenium tuckermanii (Rav. ex Mont.) Thomson
Evernia perfragilis Llano, J. Wash. Acad. Sci. 41: 199
(1951).
(see Alectoria arctica Elenk. & Savicz).
Everniastrum fragile Sipman, Proc. Kon. Ned. Akad. Wetensch.
Ser. C. 83: 348 (1980).
ISOTYPE. Colombia: Caldas, below hotel Termales, Nevado del
Ruiz. On grazed secondary shrub vegetation. H. Sipman & H.
Valencia 10512, 4 February 1979 (CANL 91358). Hale: Lich.
Amer. Exs. No. 235.
Fellhanera endopurpurea Hafellner & Vézda, Nova Hedwigia 52:
76 (1991).
ISOTYPE. Australia: New South Wales, Border Ranges National
Park, Wiangaree, Brindle Creek. Foliicola in silva
(Nothofagus moorei). J. Hafellner, P. Merotsy & R. Rogers, 30
313
August 1986 (CANL 103382). Vézda: Lich. Sel. Exs. No. 2454.
Fulgensia canariensis Follm. & Poelt, Philippia 4: 370
(1981).
ISOTYPE. Canary Islands: Tenerife, kleinfeldrig auf
lichtoffenen, weitgehend ebenen oder massig geneigten
flachgriindigen Léssinseln zwischen Hartlaubstrauchern auf
Bergriicken im Diploschistetum albescentis Klem. G. Follmann,
March 1980 (CANL 77635). Follmann: Lich. Exs. Sel. No. 345.
Glyphis achariana Tuck., Amer. J. Sci. Arts, ser. 2, 25: 429
(1858).
ISOSYNTYPE. United States: South Carolina, Santee Canal. H.W.
Ravenel, 4 February 1858 (CANL 68731). Reliq. Tuck. No. 45.
Gonohymenia cribellifera ssp. macrocarpa Henssen, Henssen:
Lich. Cyanoph. Fungi Sax. Exs. Fasc. 2: 6 (1990).
ISOTYPE. Canary Islands: Fuerteventura, Sukkulentenhalbwtiste
auf basisch-kristallinem Gestein, Passstrasse zwischen
Betancuria und Pajara nahe Vega del Rio de las Palmas; auf
sonnenexponierten Sickerwasserflachen bei 550-580 m. A.
Henssen 30731la, 31.1/1.2. 1986 (CANL 102543). Henssen: Lich.
Cyanoph. Fungi Sax. Exs. No. 37.
Gonohymenia lusitanica Henssen, Henssen: Lich. Cyanoph. Fungi
Sax. Exs. Fasc. 2: 7 :(1990).
ISOTYPE. Portugal: Baixo-Alentejo, Garique der Serra da
Arrabida; auf Sickkerwasserstreifen und in Mulden auf §S
exponierten Kalkfelsen in Meeresnahe. A. Henssen & D. Schafer
21775g, 9 August 1971 (CANL 102545). Henssen: Lich. Cyanoph
Fungi Sax. Exs. No. 39.
Gonohymenia undulata Henssen, Henssen: Lich. Cyanoph. Fungi
Sax. Exs. Fasc.,.2: 5 (1990).
ISOTYPE. Canary Islands: Fuerteventura, Sukkulentenhalbwiste
auf basisch-kristallinem Gestein, Passstrasse zwischen
Betancuria und Pajara nahe Vega del Rio de las Palmas; auf
sonnenexponierten Sickerwasserflachen bei 550-580 m. A.
Henssen 30736b, 31.1/1.2.1986 (CANL 102540). Henssen: Lich.
Cyanoph. Fungi Sax. Exs. No. 35.
PARATYPE. Canary Islands. A. Henssen 32048a (CANL 102541).
Graphina aibonitensis Fink, Mycologia 19: 215 (1927).
ISOTYPE. Puerto Rico: Aibonito. In open woods on bark. B.
Fink 2017, 1 July 1916 (CANL 2085).
Graphina muscicola Kalb, Kalb: Lich. Neotr. Fasc. 10: 8
(1988).
ISOTYPE. Ecuador: Napo, Bei Sarayacu, zwischen Tena und
Baeza. Ad einem feuchten Erdanriss, tiber Moosen. K. & A.
Kalb, 10.9.1987 (CANL 102711). Kalb: Lich. Neotr. No. 422.
Graphis subvirginalis Nyl., Bull. Torrey Bot. Club 16: 106
(1889).
ISOTYPE. United States: Florida, Fort George. On Ilex. W.W.
Calkins, 1888 (CANL 2094).
314
= Graphina subvirginalis (Nyl.) Mull. Arg.
Gyalectidium colchicum Vézda, Folia Geobot. Phytotax., Praha
18: 58 (1983).
ISOTYPE. Russia: Colchis (Transcaucasia occid.) distr.
Lazarevskoje: in faucibus rivi Dagomys Zapadnyi, supra vicum
Treja Rota. Ad folia Buxi colchicae. A. Vézda, 25 June 1979
(CANL 85979). Vézda: Lich. Sel. Exs. No. 1866.
Gyalideopsis alnicola Noble & Vézda, Folia Geobot. Phytotax.,
Bohemoslov. 14: 62 (1979).
ISOTYPES. Canada: British Columbia, Saltspring Island, road
to Mount Maxwell. On Alnus rubra. W.J. Noble 5811A, 3 (30 in
protologue) June 1976 (CANL 75390 & 75391).
Gyalideopsis kalbii Vézda, Mitt. Bot. Staatssamml. Mtinchen
19: 153 (1983).
ISOTYPES. Brazil: Sado Paulo, Ilha de SAo Sebastido, etwa 130
km 6stlich von S&ao Paulo. Osthang des Morro das Tacas; tber
Moosen, an einem Wegabstich. K. Kalb & J. Poelt, 7 July 1979
(CANL 89171 & 89439). Vézda: Lich. Sel. Exs. No. 1965; Kalb:
Lich. Neotr. No. 323.
Gyalideopsis trapperi Kalb & Vézda, Biblioth. Lichenol. 29:
49 (1988).
ISOTYPE. Ecuador: Napo, Bei Sarayacu, zwischen Tena und
Baeza. Als Erstbesiedler auf herumliegenden
Urgesteinsbloécken. K. & A. Kalb, 10.9.1987. (CANL 102712).
Kalb: Lich. Neotr. No. 423.
Gyalideopsis vezdae Kalb, Kalb: Lich. Neotr. Fasc. 6: 10
(1983).
ISOTYPE. Brazil: Mato Grosso, Serra dos Coroados, Buriti,
Naturschutzgebiet der evangelischen Schule von Buriti, etwa
6 km sltidwestlich von Buriti. In einem submontanen Regenwald.
K. Kalb, 8 July 1980 (CANL 84778). Kalb: Lich. Neotr. No.
2297
Gyrophoropsis dwaliensis R&dsdnen, Arch. Soc. Zool.-Bot. Fenn.
"Vanamo" 6: 80 (1952).
ISOTYPE. India: N. W. Himalaya, Dwali, Almora District. 8500
ft. D.D. & A.M. Awasthi, 21 May 1950 (CANL 10771). |
= Umbilicaria badia Frey
Haematomma inexpectatum Kalb, Kalb: Lich. Neotr. Fasc. 12: 11
(1991).
ISOTYPE. Mexico: Baja California Sur. Along Rte 22; 16 km W
of Ciudad Constitui6én, Southern Magdalena Region of the
Socnoran Desert, sandy coastal plain. On bark of deciduous
trees. K. & A. Kalb & T. Nash, 1 January 1991 (CANL 103777).
Kalb: Lich. Neotr. No. 506.
Haematomma pustulatum Brodo & Culb., Bryologist 89: 203
(1986).
ISOTYPE. United States: North Carolina, Orange Co.,
Hillsborough, Duke Forest (Quarry Division); near the quarry.
315
On Acer rubrum. W.L. Culberson & C.F. Culberson 19701, 22
November 1984 (CANL 96379). Vézda: Lich. Sel. Exs. No. 2152.
Hafellia fosteri Imsh. & Sheard, Bryologist 95: 85 (1992).
PARATYPE. Canada: British Columbia, Discovery Island, 3 km
east of Oak Bay, Victoria. On driftwood. A.M. Crossley & W.J.
Noble 5335a, 3 August 1975 (CANL 102584).
Heppia uruguayensis Rasdnen, Arch. Soc. Zool.-Bot. Fenn.
"Vanamo" 2: 48 (1947).
HOLOTYPE. Uruguay: Dept. PaysandtG. On rocky banks of river
Queguay near Paysandda. I.M. Lamb 3107, 10 March 1946 (CANL
983).
= Thelidium umbilicatum Th. Fr.
Heterodermia desertorum Kalb, Kalb: Lich. Neotr. Fasc. 8: 10
(1984).
ISOTYPE. Peru: Lima, Ktistenwliste bei Huacho, etwa 150 km
nérdlich von Lima. An kleinen Felsbrocken in der Nebelzone.
K. Kalb & G. Plébst, 28 July 1979 (CANL 89440). Kalb: Lich.
Neotr. °No: 324.
Heterodermia sitchensis Goward & Noble ex Goward, Bryologist
87: 366 (1984).
ISOTYPE. Canada: British Columbia, Vancouver Island, 12 km SE
of Tofino, Schooner Cove. On twigs of Picea sitchensis. T.
Goward, Kirkvold & McGrenere 83-326, 30 March 1983 (CANL
95990).
Heterothecium pachycheilum Tuck., Synops. N. Amer. Lich. 2:
56 (1888).
ISOSYNTYPE. United States: South Carolina, Santee Canal. H.W.
Ravenel, 1857 (CANL 68775). Reliq. Tuck. No. 106.
= Megalospora pachycheila (Tuck.) Sipman
Homothecium sorediosum Henssen, Bot. Not. 132: 271 (1979).
ISOTYPE. Chile: Malleco, Parque Nacional de cContulmo,
Stidbuchenwald mit Nothofagus obliqua; auf Erde und briichigem
Sandstein einer Wegbéschung. A. Henssen, G. Vobis & J. Red6én
24271, 22 February 1973 (CANL 102536). Henssen: Lich.
Cyanoph. Fungi Sax. Exs. No. 31.
Hubbsia lumbricoides W. Weber, Svensk Bot. Tidskr. 59: 61
(1965).
ISOTYPE. Mexico: Baja California, Isla Guadalupe. On seaward-
facing basalt cliffs at Melpomene Cove, southernmost tip of
the island. W.A. Weber & C.J. McCoy, 21 April 1963 (CANL
20874). Weber: Lich. Exs. Colo. No. 85.
= H. californica (Rasdnen) W. Beber
Huilia circumnigrata Vézda, Folia Geobot. Phytotax. Praha 14:
205 (1979).
ISOTYPE. Jugoslavia: Montenegro, Sinus Kotor dictus, supra
pagum Morin. Ad saxa quartzitica. A. Vézda, 18 July 1975
(CANL 73828). Vézda: Lich. Sel. Exs. No. 1665.
316
Hydrothyria venosa J. Russell, Proc. Essex Inst. 1: 188
(1853).
ISOSYNTYPE. United States: Vermont, Brattleboro. In rivulets
from Wantasquit Mt. J.L. Russell, 16 July 1851 (CANL 23259).
Relig sy LUCK eaN On 1,
Hyperphyscia mobergii Kalb, Kalb: Lich. Neotr. Fasc. 11: 9
(1990).
ISOTYPE. Venezuela: Falcon, Distr. Acosta, Zwischen Boca da
Tocuyo und San Lorenzo, etwa 20 km NW von Chichiriviche. K.
& A. Kalb, 24 August 1989 (CANL 102751). Kalb: Lich. Neotr.
No. 463.
Hypocenomyce leucococca R. Sant. in Moberg, Publ. Herb. Univ.
Uppsala, Thunbergia 2: 3 (1986).
ISOTYPE. Sweden: Harjedalen Prov., Tanndas par., c. 1 km E of
Ramundbergets Fjdllgard. On the trunk of a birch. R.
Santesson 27901, 5 August 1977 (CANL 95082). Moberg: Lich.
Sel. Exs.:Upss No.6.
Hypogymnia heterophylla Pike, Mycotaxon 16: 157 (1982).
ISOTYPE. United States: California, Mendocino Co., Pygmy
Forest Reserve, Van Damme State Park. On stunted conifers.
M.E. Hale 49365, 9 June 1977 (CANL 91360). Hale: Lich. Amer.
Exs.: No. 237...
Hypogymnia hokkaidensis Kurok., Mem. Natl. Sci. Mus. (Tokyo)
427620 (1970).
ISOTYPE. Japan: Hokkaido, Prov. Nemuro, Ochiishi. On trunk of
Picea glehnii. S. Kurokawa 65826, 2 September 1965 (CANL
45315). Kurokawa: Lich. Rar: Crit. Exs: No. 167.
Hypogymnia kiboensis Dodge, Ann. Missouri Bot. Gard. 46: 50
(1959).
ISONEOTYPE. Tanzania: Kilimanjaro, above Peter's Hut. In the
upper part of the Philippia region, on "frost-lifted ground".
O. Hedberg 1354, 23 June 1948 (CANL 99605). Moberg: Lich.
Sel. Exs. Ups. No. 75.
= Xanthoparmelia kiboensis ( Dodge) Krog & Swinscow
Hypogymnia mollis Pike & Hale, Mycotaxon 16: 161 (1982).
ISOTYPE. United States: California, San Luis Obispo Co., Los
Osos Oak Reserve, near Los Osos. On low shrubs in sandy area.
M.E. Hale 57768, 31 July 1980 (CANL 91361). Hale: Lich. Amer.
Exs. No’ 238.
Hypogymnia oceanica Goward, Bryologist 91: 229 (1988).
HOLOTYPE. Canada: British Columbia, Vancouver Island, 5 km SW
of Sutton Pass. Trunk and branches of Pinus contorta. T.
Goward 83-302, 20 March 1983 (CANL 87115).
PARATYPE. Canada: British Columbia. I. M. Brodo 18066. (CANL
69916).
Involucrothele ligurica Servit, Webbia 10: 443 (1954).
ISOTYPE. Italy: Liguria occid., Spotorno (Savona), Loco
Lajolo. Maceriicola. Sbarbaro, March 1948 (CANL 985).
317
Ionaspis ventosa P. Jgorg. & R. Sant., Nord. J. Bot. 9: 433
(1989).
ISOTYPE. Sweden: Hdrjedalen Prov., Tannds par., Mt. Réstvalen
(c. 5 km W of Mt. Stora Mittakldppen). On pebbles at top of
the mountain. R. Santesson 32679, 14 August 1989 (CANL
102824). Moberg: Lich. Sel. Exs. Ups. No. 85.
Jenmania osorioi Henssen, Lichenologist 5: 447 (1973).
ISOTYPE. Uruguay: Salto, Uruguay River, Salto Grande Island.
Auf dem oberen Teil eines grossen Steins unterhalb der
Hochwasserlinie. H. Osorio 6675, 4-6 February 1974 (CANL
102583). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 29.
Kalbiana brasiliensis Henssen, Kalb: Lich. Neotr. Fasc. 10:
9 (1988).
ISOTYPE. Brazil: Bahia, Chapada Diamantina, Serra do
Tombador, etwa 1 km vor der Stadt Morro de Chapea. Am Rand
von natiirlichen, kleinen Wasserbecken, an Sandstein. K. Kalb,
20 July 1980 (CANL 102715). Kalb: Lich. Neotr. No. 426.
Lecanactis premnea var. chloroconia Tuck., Gen. Lich. p. 194
(1872).
ISOSYNTYPE. United States: New Hampshire, Mt. Washington. E.
Tuckerman, September 1848 (CANL 68776). Reliq. Tuck. No. 107.
= L. chloroconia Tuck.
Lecanactis subdryophila Follm. & Vézda, Philippia 3: 271
(1977).
ISOTYPE. Chile: Prov. Aconcagua. Kieternpflanzung im
Windschatten der Isla Seca in der Bucht von Zapallar. G.
Follmann, January 1965 (CANL 67506). Follmann: Lich. Exs.
Sel. No. 247.
Lecania constricta W. Weber, Weber: Lich. Exs. Colo. Fasc.
P60 V3. (L984) .
ISOTYPE. United States: Colorado, Boulder Co., Outer
foothills of the Front Range, "The Flatirons", on southwest
edge of City of Boulder. On bark of Populus angustifolia just
S of mouth of Gregory Canyon. W.A. Weber, J.W. Sheard & R.A.
Anderson, 5 September 1981 (CANL 88658). Weber: Lich. Exs.
Colo. iNo. 614.
Lecania macrocarpa B. de Lesd., Bull. Soc. Bot. France 99:
146 (1952).
ISOTYPE. Italy: Gorzegno, Le Langhe (Piemonte). Sbarbaro 319,
September 1951 (CANL 12830).
Lecania sbarbaronis B. de Lesd., Bull. Soc. Bot. France 97:
170 (1950).
ISOTYPE. Italy: Varazze per Invrea. Maceriicola in via
publica. Sbarbaro, June 1949 (CANL 12836).
Lecanora aeruginascens H. Magn., Medd. Géteb. Bot. Trddg. 18:
26) 41950).
ISOTYPE. Uruguay: Montevideo, Carrasco, Cafiada de las
Cantberas. Sobre rocas expuestas al sol. H. Osorio, 17
318
January 1948 (CANL 11644).
Lecanora austrooceanica Hertel & Leuck., Hertel: Lecideaceae
Exs. Fasc. “12:2 (1990).
ISOTYPE. New Zealand: Southland. Bluff, S of Ocean Beach.
Coastal rocks. H. Hertel 30808, 12 November 1985 (CANL
102505). Hertel: Lecideaceae Exs. No. 224.
Lecanora bolanderi Tuck., Proc. Amer. Acad. Arts 6: 266
(1864).
LECTOPARATYPE. United States: California, Marin County. H.N.
Bolander (CANL 68831).
= Cladidium bolanderi (Tuck.) Ryan
Lecanora caesiorubella subsp. merrillii Imsh. & Brodo, Nova
Hedwigia 12: 28 (1966).
ISOTYPE. United States: California, Berkeley. M.A. Howe,
February 1893 (CANL 19545). Cummings: Dec. N. Amer. Lich. No.
ABS YS).
Lecanora caesiosora Poelt, Denkschr. Regensb. Bot. Ges 26: 82
(1966).
(see Lecanora soralifera H. Magn.).
Lecanora californica Brodo, Beih. Nova Hedwigia 79: 107
(1984).
HOLOTYPE. United States: California, San Mateo Co. Rocks near
the sea. M.A. Howe, February 1893 (CANL 19543). Cummings:
Dec. N. Amer. Lich. No. 131.
PARATYPES. United States: California. I.M. Brodo 20455 (CANL
76211); Oregon. I.M. Brodo (CANL 77027).
Lecanora christoi W. Weber, Bryologist 78: 206 (1975).
ISOTYPE. United States: Colorado, Garfield Co. 8 km N of
Rifle. Sandstone ridge on west side of Rifle Creek along
south edge of Grand Hogback 3.2 km SW of Rifle Gap. On talus
blocks. W.A. Weber, G. Kunkel & J. Munger, 12 June 1974 (CANL
52208). Weber: Lich. Exs. Colo. No. 458.
Lecanora cinereofusca var. appalachensis Brodo, Beih. Nova
Hedwigia 79: 116 (1984).
HOLOTYPE. United States: Arkansas, Newton Co., Lost Valley
State Park, 22 miles SW of Harrison. On sandstone block at
edge of brook. I.M. Brodo 23512, 11 August 1979 (CANL 76592).
PARATYPE. Canada: Nova Scotia. J. Macoun (CANL 11825).
Lecanora concinna ("concinnum") Thomson, Bryologist 75: 358
(1972).
ISOTYPE. United States: Alaska, north end at Franklin Bluffs
on the Sagavanirktok River. On smooth boulders on gravel
bench. J.W. Thomson, S. Shushan & J. Koranda, 1958 (CANL
44988). Thomson: Lich. Arct. No. 95.
= a species of Aspicilia
Lecanora congesta Clauz. & Vézda, Portugaliae Acta Biol.,
ser. ‘By (9% -332 (1969).
O19
PARATYPE. France: Gallia, Bouches-du-Rhéne, Marseille, insula
Maire. Ad saxa calcarea maritima, in septentrionem
spectantia. G. Clauzade, June 1968 (CANL 55614). Vézda: Lich.
Sel. Exs. No. 779.
Lecanora freyi Poelt, Mitt. Bot. Staatssamml. Miinchen 2: 464
(1958).
ISOTYPE. Austria: Bregenzer Wald, Ifenstock, Vorarlberg
(Kleines Walsertal): Kleiner Gipfel am H&ahlekopf bei etwa
2000 m, stidseitig auf Kieselkalk. J. Poelt, July 1958 (CANL
52798). Poelt: Lich. Alpium No. 73.
Lecanora fuliginosa Brodo, Beih. Nova Hedwigia 79: 124
(1984).
HOLOTYPE. Canada: Ontario, Thunder Bay District, Sibley
Peninsula, 1.3 miles east of Silver Islet. On exposed rock on
bluff. I.M. Brodo 13735, 13 June 1968 (CANL 77033).
PARATYPES. Canada: New Brunswick. S.P. Gowan 3588 (CANL
82233); Ontario. I.M. Brodo 5941 (CANL 22789), A. Henssen
14029c (CANL 77237); Quebec. I.M. Brodo 18841 (CANL 38740),
E. Lepage 16766 (CANL 34969); Northwest Territories. J.wW.
Thomson 12087 (CANL 23472).
Lecanora homalea H. Magn., Medd. Géteb. Bot. Trddg. 18: 217
(1950).
PARATYPE. Uruguay: Montevideo, Carrasco, Cafiada de las
Canteras. Sobre postes de madera. H. Osorio 1761, 17 January
1948 (CANL 12081).
Lecanora imshaugii Brodo, Beih. Nova Hedwigia 79: 137 (1984).
Holotype. Canada: Ontario, Ottawa, Rockcliffe. On a pine
trunk. J. Macoun 345, 16 April 1896 (CANL 11626).
PARATYPES. Canada: Ontario. J. Macoun 347 (CANL 12443);
United States: Delaware. I.M. Brodo 9558 (CANL 21882),
Virginia. I.M. Brodo 9435B (CANL 21780).
Lecanora insolata H. Magn., Kongl. Svenska Vetenskapsakad.
Hand tge1 727-137 3(1939)..
ISOTYPE. Sweden: Vdastergétland, G6dteborg, Delsj6karr. On
steep, sunny rocks in the heath. A.H. Magnusson, 11-10-1937
(CANL 20146). Magnusson: Lich. Sel. Scand. Exs. No. 323.
Lecanora kutakii Senft, Preslia 4: 26 (1926).
ISOTYPE. Czechoslovakia: Sudeti occid., KrkonoSe, in valle
Obfi dil loco dicto Kiesberg. V. Kuf&k, 1921 (CANL 33384).
= L. gisleri Poelt & Ullrich
Lecanora luteovernalis Brodo, Bryologist 84: 521 (1981).
HOLOTYPE. Canada: Northwest Territories, Dist. of Franklin,
Bathurst Island, Polar Bear Pass area. I.M. Brodo & R.R.
Ireland 19435, 23 July 1973 (CANL 59491).
PARATYPES. Canada: Yukon. G.W. & G.G. Douglas 8895 (CANL
56763); G.W. Scotter 19127, 19252, 19257, 19487, 19525, 19530
& 19531 (CANL 50183, 49740, 49739, 54313, 54315, 54316 &
54317); Northwest Territories. P. Barrett 0807. (CANL 34299);
G.R. Brassard 4253B (CANL 28932); I.M. Brodo 19388 (CANL
320
53565); S.D. MacDonald (CANL 53033); G.R. Parker SP-70-184C
(CANL 33112); R.M. Schuster 35941 (CANL 5572); G.W. Scotter
18487 (CANL 70135); J.W. Thomson 8689 (CANL 45361).
Lecanora mellea W. Weber, Bryologist 78: 208 (1975).
ISOTYPE. United States: California, Sacramento Co., SE of
Sacramento between Slough House and Bridge House. On sloping
faces of rock outcrops. W.A. Weber & S. Kipp, 24 March 1974
(CANL 52212). Weber: Lich. Exs. Colo. No. 462.
Lecanora microbola Lamb, Rep. (Annual) Natl. Mus. Canada,
1952-53. Bull. 132: 291 (1954).
HOLOTYPE. Canada: Nova Scotia, Cape Breton Co., Cape Breton
Island, Louisbourg. On large stone near seashore. I.M. Lamb
6897, 19 July 1952 (CANL 12151).
Lecanora oleae ("Yolea") Reich. & Galun, Bull. Res. Council
Israel Sect. D, Bot. 6D: 238 (1958).
SYNTYPE. Israel: Thalpioth (Jerusalem). On Olea europea. I.
Reichert 8(1), 16 October 1933 (CANL 32294).
Lecanora opiniconensis Brodo, Mycotaxon 26: 309 (1986).
HOLOTYPE. Canada: Ontario, Leeds Co., Snake Island in Lake
Opinicon. Queen's University Biological Station, Chaffeys
Locks. I.M. Brodo 25117, 2 February 1985 (CANL 92112).
PARATYPES. Canada: Ontario. I.M. Brodo 456, 13575B, 20277,
23432B & 23543 (CANL 23295, 79030, 53929, 72543 & 80250);
R.F. Cain 26672 (CANL 44587); H.L. Dickson 154 (CANL 57666) ;
C.E. Garton 5467, 6089 & 22268 (CANL 12295, 12304 & 87731);
C.M. Wetmore 28714 & 29679 (CANL 85169 & 85259). Quebec. I.M.
Brodo 21015 (CANL 54173). United States: Maine. S. Selva 990
(CANL 98807). Michigan. C.M. Wetmore 47222 (CANL 90696).
Minnesota. C.M. Wetmore 34532 & 35014 (CANL 75807 & 75810).
S. Dakota. C.M. Wetmore 8760 (CANL 12150). Wisconsin. T.S. &
B.A. Cochrane 5601 (CANL 45366).
Lecanora orae-frigidae R. Sant., Lich. Sweden Norway, p.145
(1984).
(see Lecidea sorediata Lynge).
Lecanora pachysoma Ryan & Poelt, Bryologist 92: 518 (1989).
PARATYPES. Mexico. T.H. Nash 8804. (CANL 52389); W.A. Weber
& C.J. McCoy. (CANL 20876). Weber: Lich. Exs. Colo. No. 87.
Lecanora perplexa Brodo, Beih. Nova Hedwigia 79: 148 (1984).
HOLOTYPE. Canada: Quebec, Gatineau Co., Gatineau Park, King
Mountain area off Mountain Road, 3 mi. NW of Notch Road
intersection. On limestone cliff. I.M. Brodo 21192, 6 October
1975 (CANL 67214).
PARATYPES. Canada: Ontario. I.M. Brodo 6070 (CANL 22769), A.
Henssen 14276 (CANL 77235), J. Macoun? (CANL 11624); Quebec.
I.M. Brodo 23585 (CANL 73901); United States: Arkansas. I.M.
Brodo 13504 (CANL 76596).
Lecanora pseudopinguis W. Weber, Mycotaxon 13: 102 (1981).
ISOTYPE. Ecuador: Galapagos Islands, Isla Santa Cruz. On
321
rocks just above high tide. W.A. Weber & J. Lanier, 10 April
1976 (CANL 58215). Weber: Lich. Exs. Colo. No. 500.
Lecanora pycnocarpa H. Magn., Bot. Not. 1939: 310 (1939).
ISOTYPE. Sweden: Uppland, Vallentuna, Abydal. On Populus
tremula. A.H. Magnusson, 12 June 1934 (CANL 20132).
Magnusson: Lich. Sel. Scand. Exs. No. 305.
Lecanora salicicola H. Magn., Bot. Not. 1939: 311 (1939).
ISOTYPE. Sweden: Jamtland, Are, Skalstugan. On branches of
dying Salix. C. Stenholm, 28 July 1933 (CANL 20133).
Magnusson: Lich. Sel. Scand. Exs. No. 306.
Lecanora soralifera H. Magn., Bot. Not. 1937: 135 (1937).
ISOTYPE. Sweden: Vastergétland, Partille, northwest of
Tultered. On stone fence. A.H. Magnusson, 27 August 1936
(CANL 20111). Magnusson: Lich. Sel. Scand. Exs. No. 270.
= L. caesiosora Poelt
Lecanora superfluens H. Magn., Acta Horti Gothob. 19: 46
(1952).
ISOTYPE. Canada: Northwest Territories, Baffin Island. Head
of Clyde Fiord. On wet soil near large boulders. M.E. Hale
416, 26 June 1950 (CANL 12472).
= L. geophila (Th. Fr.) Poelt
Lecanora texana W. Weber, Mycotaxon 13: 103 (1981).
ISOTYPE. United States: Texas, Gillespie Co. (not "Llano Co.
-as given on the label), Texas Hill Country, trail to Balanced
Rock, 6.4 km N of Fredericksberg. On vertical N-facing of
large blocks. W.A. Weber, 29 April 1974 (CANL 52201). Weber:
Rich..Exs.,.Colo.iNo. 451.
Lecanora thallophila H. Magn., Bot. Not. 107: 196 (1954).
ISOTYPE. United States: Utah, San Juan Co., Devil's Canyon.
On Dermatocarpon miniatum on dry exposed sandstone. S.
Flowers 476, 21 June 1952 (CANL 71524).
Lecanora utahensis H. Magn., Acta Horti Gothob. 19: 39
(1952).
ISOTYPE. United States: Utah, Wayne Co., Ekker's Ranch. On
dry exposed sandstone. S. Flowers 3107, 20 May 1951 (CANL
12503)%%
Lecanora weberi Ryan, Mycotaxon 36: 10 (1989).
ISOTYPE. United States: Colorado, Boulder Co., Boulder
Mountain Parks, just S of city of Boulder. On scattered low
fine-grained sandstone or quartzite boulders. W.A. Weber, 10
August 1986 (CANL 97984). Weber: Lich. Colo. Exs. No. 685.
Lecidea baffiniana H. Magn., Acta Horti Gothob. 19: 43
(1952).
ISOTYPE. Canada: Northwest Territories, Baffin Island, Cape
Searle. On phonolite at the bird cliffs. M.E. Hale 391, 17-18
August 1950 (CANL 4786).
322
Lecidea carbonoidea Thomson, Bryologist 75: 356 (1972).
ISOTYPE. United States: Northern Alaska, along the Pitmegea
River, 15 miles upstream from Cape Sabine. On top of ridge.
J.W. Thomson, 10-17 July 1958 (CANL 44987). Thomson: Lich.
Arct. No. 94.
= Immersaria carbonoidea (Thomson) Esnault & Roux
Lecidea grassiana H. Magn., Medd. Gdéteborgs Bot. Trddg. 17:
60 (1947).
ISOTYPE. Argentina: Prov. Salta, Quebrada de San Lorenzo. M.
Digilio-Grassi 336, 1946 (CANL 5061).
Lecidea gypsicola Llimona, Vézda: Lich. Sel. Exs. Fasc. 47:
She GLI LES) is
ISOTYPE. Spain: Las Bardenas, Caparosso. Ad gypsum compactum
locis septentr. vel occident. spectantibus. X. Llimona, 24
September 1972 (CANL 77198). Hertel: Lecideaceae Exs. No. 45.
Lecidea (Psora) hyotheja Lamb, Farlowia 4: 444 (1955).
HOLOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Verde near
Futalaufquen, On soil in crevices of an almost vertical rock
on the shore of the lake. I.M. Lamb 5870, 1 February 1950
(CANL 5072).
PARATYPE. Argentina: Lago Cisnes. I.M. Lamb 5932 (CANL 5071).
Lecidea leproloma R. Sant., Lich. Sel. Exs. Upsal.,
Thunbergia 2: 6 (1986).
ISOTYPE. Sweden: Hadrjedalen Prov., Tann&s par., Mt Gruvvalen,
0.2 km SSW of Lake Glimsj6n (c. 1.8 km NE of Ramundbergets
Fjallgard). On a vertical rock wall. R. Santesson 31390, 19
August 1985 (CANL 95090). Moberg: Lich. Sel. Exs. Ups. No.
14.
Lecidea patagonica Lamb, Farlowia 4: 440 (1955).
HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Pathway from
Lago Frias to the Frias Glacier. On a stone beside the path
in the forest. I.M. Lamb 6029, 13 November 1950. (CANL 5290).
PARATYPE. Argentina: Lake Nahuel Huapi. I.M. Lamb 5818 (CANL
5289).
Lecidea santensis Tuck., Amer J. Sci. Arts ser. 2, 25: 428
(1858).
SYNTYPE. United States: South Carolina. H.W. Ravenel (CANL
23244)|. Relig. “Tuck. No. 15.
= Phyllopsora corallina (Eschw.) MUll. Arg.
Lecidea shushanii Thomson, Bryologist 75: 356 (1972).
ISOTYPE. United States: Northern Alaska, valley of the
Okpilak River at Okpilak Lake near Mt. Michelson. On huge
boulder at top of ridge. J.W. Thomson & S. Shushan 9316, 3
August 1958 (CANL 44982).
Lecidea sorediata Lynge, Novaya Zemlya 43: 117 (1928).
ISOLECTOTYPE. Russia: Novaya Zemlya, Nordre Kristovii holmen.
B. Lynge, 14 August 1921. (CANL 5430).
= Lecanora orae-frigidae R. Sant.
323
Lecidea ullrichii Hertel, Hertel: Lecideaceae Exs. Fasc. 10:
4 (1988).
ISOTYPE. Germany: Niedersachsen, Harz Ramelsberg bei Goslar.
H. Ullrich, 26 October 1982 (CANL 96965). Hertel: Lecideaceae
Exs. No. 190.
Lempholemma dispansum H. Magn., Bot. Not. 1939: 302 (1939).
ISOTYPE. Sweden: Dalsland, Backe, Karud. On sunny, slightly
irrigated, calcareous rock. S. Bergstrom & A.H. Magnusson, 24
June 1938 (CANL 20128). Magnusson: Lich. Sel. Scand. Exs. No.
298.
Leproloma diffusum var. chrysodetoides Laundon, Lichenologist
21: 18 (1989).
ISOTYPE. Sweden: Vdarmland Prov., Burtjdarn par., on the
western slope of the hill between Irettondetjdrnen and
Svarttjdrnen. Over mosses on perpendicular rock. L.E. Muhr
598, 8 November 1977 (CANL 99591). Moberg: Lich. Sel. Exs.
Ups. No. 61.
Leptogium caesiellum Tuck., Synopsis N. Amer. Lich. 1: 156
(1882).
ISOTYPE. United States: Illinois, Athens. EF. Hall, 1876 (CANL
23261). eReliq .etTucka No1w53.%
= L. byssinum (Hoffm.) Zwackh ex Nyl.
Letharia togashii Asah., J. Jap. Bot. 27: 295 (1952).
ISOTYPE. Japan: Honshu. Prov. Kai, Oshino-mura, Minami-Tsuru-
gun. On twigs of Picea polita. M. Togashi, 11 August 1952
(CANL 26404). Kurokawa: Lich. Rar. Crit. Exs. No. 66.
Letharia wandelensis Hue, Expedit. Antarct. Fran¢. 1903-1905,
P7568 61908 ),.
ISOTYPE. Antarctica: Graham Land (Palmer Peninsula), Booth
(Wandel) Island. Expedit. Antarct. Fran¢. 1903-1905, No. 277
& 299 (CANL 16959).
Lichen candelarius L., Spec. Plant 2: 1141, 1753.
ISONEOTYPE. Sweden: Oland Prov., Boda par., Mensalvaret. On
the top of granitic boulders. R. Santesson & T. Tonsberg
30905, 3 August 1983 (CANL 95101). Moberg: Lich. Sel. Exs.
Ups. No 25.
= Xanthoria candelaria (L.) Th. Fr.
Lichen salazinus Bory, Voyage Quatre Iles d'Afrique 3: 106
(1804).
ISOLECTOTYPE. Africa: Réunion (Bourbon) Island. Bory de St.
Vincent (CANL 75659).
= Stereocaulon salazinum (Bory) Fée
Lichenoconium parasiticum D. Hawksw., Persoonia 9: 178
(1975 \e
ISOTYPE. Denia: Sjaellandia, Herstedvester. In apotheciis
Lecanorae conizaeoidis. M.S. Christiansen 11707, 20 April
(July in protologue) 1944 (CANL 69519). Vézda: Lich. Sel.
Exs. No. 1600.
324
Lichenostigma maureri Hafellner, Herzogia 6 : 299 (1982).
PARATYPE. Italy: Siidtirol, Mendelgebirge SW von Bozen, Mte.
Roén, Larchen-Fichtenwald auf der N-Seite. J. Hafellner,
October 1976 (CANL 91566). Poelt: Pl. Graecenses No. 319.
Lichina macrospora Henssen, Biidel & Wessels, Mycotaxon 22:
171 (1985).
ISOTYPE. South Africa: Ost-Transvaal, Graskop 2430DD,
Panorama Falls, Lookout Point. Auf Sandsteinplatten am Rande
von Rinnsalen. A. Henssen & D. Wessels 28405a, 8 June 1982
(CANL 102537). Henssen: Lich. Cyanoph. Fungi sax. Exs. No.
B2e
Lichina polycarpa Henssen, Lichenologist 5: 450 (1973).
PARATYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel
Huapi, Isla Victoria. On semi-decomposed rock at side of
lake. I.M. Lamb 5822 (=5821), 26 January 1950 (CANL 2358).
Lichina rosulans Henssen, Lichenologist 4: 95 (1969).
ISOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel
Huapi, Isla Victoria. On rocks at shore of lake. I.M. Lamb
5847 (=5857), 28 January 1950. (CANL 27353)
Lobaria fulva Hoffm., Deutschl. Flora, p. 159, 1796.
ISONEOTYPE. Germany: Bavaria, Berchtesgadener Land,
Untersch6nau, Astenreitweg-Zulehenweg. Ad truncum Aceris
pseudoplatani. R. Ttirk & H. Wunder, 30 April 1991 (CANL
103428). Vézda: Lich. Sel. Exs. No. 2500.
= Xanthoria fulva (Hoff.) Poelt & Petutschnig
Lobaria fuscotomentosa Yoshim., J. Hattori Bot. Lab. 34: 311
(1971).
ISOTYPE. Japan: Honshu, Prov. Echigo. On trail between the
summit and Akayu, Mt. Naeba. On trunk of trees. S. Kurokawa
57194, 17 August 1957 (CANL 45320). Kurokawa: Lich. Rar.
CYTG. EXs.-NOvelL 7 2.
Lobaria gyrophorica Yoshim., J. Hattori Bot. Lab. 34: 275
(1971).
ISOTYPE. Taiwan: Prov. Ilan, Piyanan Pass. On trees. S.
Kurokawa 794, 16 January 1964 (CANL 68480). Kurokawa: Lich.
Rare Crit. albxa.NOw 365"
Lobaria tuberculata Yoshim., J. Hattori Bot. Lab. 34: 280
(1971).
ISOTYPE. Japan: Hokkaido, Prov. Nemuro, Ochiishi. On trunk of
Salix sp. S. Kurokawa 65767, 1 September 1965 (CANL 45326).
Kurokawa: Lich. Rar. Crit. Exs. No 178.
Maronella laricina M. Steiner, Osterr. Bot. Z. 106: 441
(1959).
ISOTYPE. Austria: Wipptal, Tirol, Steinach-Mauern. Fussteil,
NW-Flanke von Larix decidua an einem Rinnsal in Fettwiesen.
M. Steiner, August 1958 (CANL 52819). Poelt & Steiner: Lich.
Alpium No. 102.
325
Mazosia pilosa Kalb & Vézda, Folia Geobot. Phytotax., Praha
23: 203 (1988).
ISOTYPES. Brazil: Amazonas, Manaus, in ripa fluminis Rio
Negro, 100-200 km supra Manaus. Foliicola in pluviisilva. K.
Kalb, 14 October 1980 (CANL 97329 & 102720). Vézda: Lich.
Sel. Exs. No. 2227; Kalb: Lich. Neotr. No. 431.
Mazosia pseudobambusae Kalb & Vézda, Folia Geobot. Phytotax.,
Praha 23: 207 (1988).
ISOTYPE. Brazil: Sdo Paulo, Ilha de Sdo Sebastiao, 130 km ad
orientem a Sdo Paulo. Folliicola in pluviisilva. K. Kalb, J.
Poelt & H. Sipman, 6 July 1980 (CANL 97330). Vézda: Lich.
Sel. Exs. No. 2228.
Megalospora foersteri Kalb, Kalb: Lich. Neotr. Fasc. 11: 10
(1990).
ISOTYPE. Venezuela: Merida, Distr. Libertador, Pico Espejo,
SE von Mérida, Loma Redonda. Uber feuchten Moosen auf
Lichtungen eines kleinen Restwdldchens. K. & A. Kalb, 10
August 1989 (CANL 102753). Kalb: Lich. Neotr. No. 465.
Melanaria macounii Lamb, Rep. (Annual) Natl. Mus. Canada,
1952-53 BULL 1323328 6n41954) ¢
HOLOTYPE. Canada: Nova Scotia, Victoria Co., Cape Breton
Island, Franey Mountain trail near Ingonish. On trunk of
Fagus. I.M. Lamb 6985, 6 August 1952 (CANL 11613).
= Pertusaria macounii (Lamb) Dibben
Melanaria macounii var. meizotoca Lamb, Rep. (Annual) Natl.
Mus eCanada,.2952=53' abu) Lay 6132 2628 a Glo54)
HOLOTYPE. Canada: Nova Scotia, Cape Breton Island, Baddeck.
JseMacoun, 1953) p.pw.W,).9) July) 1898 (CANL' {11479),.
= Pertusaria macounii (Lamb) Dibben
Melaspilea fuscolimitata Fink, Mycologia 19: 211 (1927).
ISOTYPE. Puerto Rico: Yauco, Dry Hill. On bark. B. Fink 1480,
29 December 1915 (CANL 1974).
Micarea muhrii Coppins, Bull. Brit. Mus. (Nat. Hist.), Bot.
11(2): 160 (1983).
ISOTYPE. Sweden: Vermlandia, par. Lungsund, Punbd&dcken. Loco
aperto ad truncum decorticatum supra rivum. L.E. Muhr 285
(2851 in protologue), 15 July 1980 (CANL 89160). Vézda: Lich.
Sel. Exs. No. 1954.
= Micaria vulpinaris (Nyl.) Muhr
Microthelia atramentea Norman, Bot. Not. 1867: 88 (1867).
ISOTYPE. Norway: Finmark, Tanafjord, Lille Lerpollen. J.M.
Norman, 1866 (CANL 93368).
Muhria urceolata P. Jgrg., Notes Roy. Bot. Gard. Edinburgh
44: 583 (1987).
ISOTYPE. Sweden: Vdrmland. Dalby, Gravbdcken, 6 km N of
Syssleback. L.E. Muhr 4208, 6 August 1981 (CANL 95302).
Mycoglaena quercicola R. Harris, Michigan Bot. 12: 30 (1973).
326
ISOTYPE. United States: Iowa, Bremer Co., Iowa. B. Fink, 1895
(CANL 1266).
Nephroma chubutense Lamb, Farlowia 4: 438 (1955).
HOLOTYPE. Argentina: Patagonia, Prov. Chubut, W end of Lago
Menéndez in the lower reaches of the Torrecillas stream. On
trunk of a small tree. I.M. Lamb 5916, 4 February 1950 (CANL
TOZ25 7)es
Nephroma kuehnemannii Lamb, Farlowia 4: 436 (1955).
ISOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Menéndez,
Alerzal del Norte. O. Ktihnemann 5485, 1941 (CANL 3751).
Nephroma lepidophyllum f. hypomelaenum ("hypomelaena")
Rasdnen, Farlowia 4: 439 (1955).
ISOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lago Frias,
pathway alongside the Frias stream on trunk of Nothofagus
pumilio. I.M. Lamb 6010, 12 February 1950 (CANL 3785).
Nephroma occultum Wetm., Bryologist 83: 243, !980.
ISOTYPE. United States: Oregon, Lane Co., 7 miles NE of Blue
River, H.J. Andrews Experimental Forest, 2 miles NE of the
junction of Blue River and Lookout Creek. On old growth
Pseudotsuga menziesii branches. S. Sundberg 120, 19 February
1978 (CANL 75860).
Neuropogon rohmederi Lamb, Lilloa 14: 158 (1948).
ISOTYPE. Argentina: Chubut, Lago Futalaufquén. G. Rohmeder
(Herb. Crypt. Inst. Lillo no. 2275), 1945. (CANL 17205).
= Usnea perpusilla (Lamb) F. J. Walker
Neuropogon rohmederi f. ushuaiensis Lamb, Lilloa 14: 160
(1948).
ISOTYPE. Argentina: Tierra del Fuego, Sierra Alvear, ca. 20
km ENE of Ushuaia. On rocks. R. Santesson 640a p.p., 1940
(CANL 17209).
= Usnea perpusilla (Lamb) F. J. Walker
Neuropogon trachycarpus f. elatior Lamb, Lilloa 14: 157
(1948).
ISOTYPE. Argentina: Tierra del Fuego, Sierra Alvear, southern
slope, above Las Cotorras (about 20 km ENE of Ushuaia). On
rocks in the alpine region. R. Santesson 640b, 1940 (CANL
17258).
= Usnea trachycarpa (Stirt.) MUll. Arg.
Ochrolechia gowardii Brodo, Canad. J. Bot. 69: 746 (1991).
HOLOTYPE. Canada: British Columbia, Wells Gray Park, Wavy Mt.
near Murtle Lake. On branch of Abies lasiocarpa. T. Goward
86-39-5, 13 August 1980 (CANL 96072).
Ochrolechia juvenalis Brodo, Canad. J. Bot. 69: 749 (1991).
HOLOTYPE. United States: California, Mendocino Co., Fort
Bragg. On twigs of Pinus muricata. I.M. Brodo 25301, 13 July
1986 (CANL 94666).
327
Ochrolechia montana Brodo, Canad. J. Bot. 69: 753 (1991).
HOLOTYPE. United States: Montana, Lake Co., near Soup Creek,
Swan Range. On bark. B. McCune 8854, 26 August 1877 (CANL
77559).
Ochrolechia pseudopallescens Brodo, Canad. J. Bot. 69: 755
(1991).
HOLOTYPE. Canada: Quebec, Gatineau Co., near Eardley in Picea
mariana bog between Ramsay Lake and Lac Hawley. On Picea
mariana twigs. I.M. Brodo 16280, 2 October 1969 (CANL 34613).
Ochrolechia subisidiata Brodo, Canad. J. Bot. 69: 759 (1991).
HOLOTYPE. United States: Arizona, Gila Co., McFadden Peak, 15
mi. S of Young. On dead Juniperus branch. T.H. Nash 11166
(CANL 58113).
Ochrolechia trochophora var. pruinirosella Brodo, Canad. J.
Bot. 69: 763 (1991).
PARATYPES. Canada: New Brunswick. Gowan 3129 (CANL 81779);
Ontario. I.M. Brodo 14680 & 16130 (CANL 27752 & 60142);
Quebec, I.M. Brodo 9630, 16895 & 25290 (CANL 21653, 72040 &
93690); Prince Edward Island. J. Fabiszewski 7C & 9A (CANL
9329984933 00)«:
Ochrolechia tuckermanii Vers., Beih. Nova Hedwigia 1: 97
(1962).
ISOTYPE. United States: Massachusetts, Manchester. E.
Tuckerman, 1839. (CANL 23277). Reliq. Tuck. No. 114 ("No. 14"
in protologue).
= O. yasudae Vainio
Omphalodina bullata Follm. & Crespo, Philippia 3: 24 (1976).
ISOTYPE. Spain: Prov. Toledo, Koloniebildend an
halbschattigen, luftfeuchten, teils lberhangenden,
verhdltnismassig glatten Silikatfeisen. G. Follmann & A.
Crespo, 1975. (CANL 58316). Follmann: Lich. Exs. Sel. No.
174.
= Rhizoplaca bullata (Follm. & Crespo) Leuck. & Poelt.
Opegrapha albidoatra Fink, Mycologia 19: 210 (1927).
PARATYPE. Puerto Rico: Naranjito. In woods on bark. B. Fink
298, 27 November 1915 (CANL 1903).
Opegrapha astraea Tuck., Lich. Calif. p. 33 (1866).
SYNTYPES. United States: South Carolina, Santee Canal. On
Acer rubrum. H.W. Ravenel 466, 472 (CANL 68781, 68782).
Reliq. Tuck. Nos. 119a, 119b.
Opegrapha pertusariicola Coppins & P. James, Lichenologist
11: 164'-(1979).
ISOTYPE. Great Britain (England): Caledonia, Mid-Perthshire,
Aberfeldy, Glen Lyon, prope Woodend. In thallo Pertusariae
leucoplacae vigens, ad corticem Coryli. P. James & B.J.
Coppins 3834, 11 September 1976 (CANL 73861). Vézda: Lich.
Sel. Exs. No. 1699.
328
Opegrapha subabnormis Fink, Mycologia 19: 210 (1927).
ISOTYPE. Puerto Rico: Mayaguez. In open field on bark. B.
Fink 1245, 33 December 1915 (CANL 1946).
Oropogon colibor Essl., Syst. Bot. Monogr. 28: 60 (1989).
ISOTYPE. Costa Rica: Cartago Prov., about 20.4 km SE of El
Palme. On tree bark. F. Almeda 2143B, 27 December 1973 (CANL
98282).
Oropogon diffractaicus Essl., Syst. Bot. Monogr. 28: 61
(1989).
ISOTYPE. West Indies: Dominican Republic, Sierra de la
Pelonia, Maciso Central, Cordillera Central. On small shrub
in open. C.M. Wetmore & H.A. Imshaug 3652, 9 August 1958
(CANL 16620).
Oropogon mexicanus Essl., Syst. Bot. Monogr. 28: 92 (1989).
ISOTYPE. Mexico: State of Oaxaca, Trail leading to Cerro San
Felipe. On conifers. M.E. Hale 20706, 1 April 1960 (CANL
31064).
Pannaria leucostictoides Ohlsson, Bryologist 76: 379 (1973).
ISOTYPE. Canada: British Columbia, Queen Charlotte Islands,
Moresby Island, Skidegate Lake. On branches of Malus. I.M.
Brodo 11085, 9 July 1967 (CANL 29174).
Parmelia alabamensis Hale & McCull., Bryologist 71: 44
(1968).
ISOTYPE. United States: Alabama, St. Clair Co., 6 miles E of
Leeds. On sandstone outcrops. M.E. Hale 24072, June 1966
(CANL 31055).
= Paraparmelia alabamensis (Hale & McCull.) Elix & Johnston
Parmelia albertana Ahti, Bryologist 72: 236 (1969).
ISOTYPE. Canada: Alberta, near SW tip of Big Lake, ca. 10 mi.
NW of Edmonton City. On Populus balsamifera. T. ‘Ahti LItAas
7 July 1967 (CANL 28560).
= Melanelia albertana (Ahti) Essl.
Parmelia ambigua var. halei Tuck., Synops. N. Amer. Lich. 1:
66 (1882).
ISOSYNTYPE. United States: South Carolina. On Pinus mitis.
H.W. Ravenel (CANL 68744). Reliq. Tuck. No. 65.
= Parmeliopsis subambigua Gyelnik
Parmelia chlorochroa Tuck., Proc. Amer. Acad. Arts 4: 383
(1860).
ISOSYNTYPE. United States: North Dakota, Missouri River. F.V.
Hayden, 1859-60 (CANL 68784). Reliq. Tuck. No. 121.
= Xanthoparmelia chlorochroa (Tuck.) Hale
Parmelia crinita f. varians G. K. Merrill, Bryologist 11: 95
(1908).
HOLOTYPE. Canada: Ontario, Carleton Place. On elm trees ina
Swamp. J. Macoun 3828, 12 May 1900 (CANL 13348).
= Parmotrema subtinctorium (Zahlbr.) Hale
329
Parmelia cryptoxanthoides Kurok., Bull. Natl. Sci. Mus.
Tokyo. 17: 297 (1974).
ISOTYPE. Brazil: Parana, Jardim Paraizo, 9 km west of
Curitiba. On trees. S. Kurokawa 8245, 9 July 1971 (CANL
57029). Kurokawa: Lich. Rar. Crit. Exs. No. 232.
Parmelia elabens Kurok., Bull. Natl. Sci. Mus. Tokyo. 17: 298
(1974).
ISOTYPE. Brazil: Parana, Jardim Paraizo, 9 km west of
Curitiba. On trees. S. Kurokawa 8253, 9 July 1971 (CANL
57030). Kurokawa: Lich. Rar. Crit. Exs. No. 233.
Parmelia dubia var. scrobiculata B. de Lesd., Bull. Soc. Bot.
BVance 499: 145941952) :.
ISOTYPE. Italy: Liguria sabatia, Spotorno, tinicola loco
"Collina". Sbarbaro, 19 December 1951 (CANL 13700).
= Punctelia perreticulata (Rdsdnen) Wilhelm & Ladd
Parmelia frondifera G. K. Merrill, Bryologist 11: 91 (1908).
HOLOTYPE. Canada: Ontario, Essex Co., Point Pelee. On cedar
bark. J. Macoun 3627, 27 May 1901 (CANL 13193).
= Punctelia bolliana (Miill. Arg.) Krog
Parmelia hygrophila Goward & Ahti, Ann. Bot. Fenn. 20: 9
(1983).
ISOTYPE. Canada: British Columbia, Kokanee Creek Provincial
Park, 17 km east of Nelson. On trunk of Betula papyrifera. T.
Goward 81-1601, 25 July 1981 (CANL 86317).
Parmelia gloriosa Kurok., Stud. Cryptog. Papua New Guinea p.
135 (1979).
ISOTYPE. Papua New Guinea: Central District, Mt. Albert
Adward, around Abios Hut. On bark of Pandanus sp. S. Kurokawa
9113, 31 October 1975 (CANL 68483). Kurokawa: Lich. Rar.
Crit Exs.)No,.°'368.
Parmelia omphalodes subsp. glacialis Skult, Ann. Bot. Fenn.
Zee 2cuLe (1985)\,
HOLOTYPE. Canada: Northwest Territories, Dist. of Franklin,
Prince Patrick Island, Mould Bay. On damp polygon soil. S.D.
MacDonald, 24 July 1952 (CANL 13656).
PARATYPES. Canada: Northwest Territories. L. C. Bliss 2.
(CANL 70510); M. Kuc 6, AA-14, AK-11, AK-12, AK-13, AM-24,
AN-15, LC-13 & LD-11 (CANL 29811, 29854, 29871, 29818, 29787,
29850, 29840, 29774 & 29779); S.D. MacDonald (CANL 13647);
D.B.O. Savile 4102 (CANL 46004); D.W. Smith (CANL 63481).
= Parmelia skultii Hale
Parmelia pacifica Kurok., Mem. Natl. Sci. Mus. Tokyo 11: 28
(1978).
ISOTYPE. Japan: Bonin Islands, Hahajima Island, En route
between Oki-mura and Cape Minami-zaki. On bark of Pandanus
boninensis. S. Kurokawa 77014, 3 August 1977 (CANL 68485).
Kurokawa: Lich. Rar. Crit. Exs. No. 370.
Parmelia physodes var. enteromorpha f. rugosa G. K. Merrill,
330
Bryologist 11: 86 (1908).
HOLOTYPE. Canada: British Columbia, near MacLeodes Lake. J.
Macoun 696, 27 June 1875 (CANL 14331).
= Hypogymnia rugosa (G. K. Merrill) Pike
Parmelia physodes f. subisidioides G. K. Merrill, Bryologist
Dis 86m (1908).
SYNTYPE. Canada: Alberta. On old logs at Laggan. J. Macoun
3635, 26 June 1904 (CANL 14298).
= Hypogymnia austerodes (Nyl.) Rdasdnen
Parmelia ralla Brusse, Mycotaxon 27: 241 (1986).
ISOTYPE. South Africa: Swartberg range, Seven Weeks poort,
8.6 km NW of Amalienstein. On Table Mountain sandstone
boulders. F. Brusse 4935, 10 February 1986 (CANL 97358).
Parmelia reptans Kurok., Austral. J. Bot. 21: 137 (1973).
ISOTYPE. Australia: Victoria, 6 miles west of Red Cliffs
along the Werrimull Road. On soil. S. Kurokawa 6621, 8
December 1965 (CANL 68487). Kurokawa: Lich Rar. Crit. Exs.
NOSUE3 7 25
= Xanthoparmelia reptans (Kurok.) Elix & Johnston
Parmelia saximontana R. Anderson & W. Weber, Bryologist 65:
23081962) e.
ISOTYPE. United States: Colorado, Boulder Co., Dakota Group
just NW of Boulder. On sandstone. R.A. Anderson & W.A. Weber
L-30072, 6 April 1962 (CANL 20831). Weber: Lich. Exs. Colo.
Now (41.
= Melanelia tominii (Oxner) Essl.
Parmelia semansiana Culb. & C. Culb., Mycologia 72: 128
(1980).
ISOTYPE. United States: Arkansas, Polk Co., 5 km ad
septentriones versus a Mena, in loco dicto Blue Haze Vista
prope viam dictam Talimena Skyline Drive. Ad saxa in silva.
W.L. Culberson & C.F. Culberson 16074, 20 August 1972 (CANL
76386). Vézda: Lich Sel. Exs. No. 1737.
= Punctelia semansiana (Culb. & C. Culb.) Krog
Parmelia spinibarbis Kurok., Bull. Natl. Sci. Mus. Tokyo. 17:
299 8( 197A).
ISOTYPE. Brazil: Rio de Janeiro, Petrépolis. On trees. S.
Kurokawa 8348, 17 July 1971 (CANL 57035). Kurokawa: Lich.
Rar. Crity Exs sj) Now 2387
Parmelia squarrosa Hale, Phytologia 22: 29 (1971).
ISOTYPE. United States: Virginia, Madison Co., Shenandoah
National Park. On Quercus alba. M.E. Hale 36949, 30 April
1979 (CANL 38279). Hale: Lich. Amer. Exs. No. 142.
Parmelia subdistorta Kuroki 2) Hattorr Bot... bab: 32-7 ie
(1969).
ISOTYPE. Australia: Victoria, 6 miles west of Red Cliffs
along the Werrimull Road. On soil. S. Kurokawa 6617, 8
December 1965 (CANL 45332). Kurokawa: Lich. Rar. Crit. Exs.
331
No.4, 184.
= Xanthoparmelia subdistorta (Kurok.) Hale
Parmelia sulcifera Kurok., Bull. Natl. Sci. Mus. Tokyo. Ser.
Bees: 371982)
ISOTYPE. Australia: New South Wales, 9 miles E of Cooma on
Numeralla Road. On rocks. S. Kurokawa 6450, 2 December 1965
(CANL 87340). Kurokawa: Lich. Rar. Crit. Exs. No. 583.
= Xanthoparmelia sulcifera (Kurok.) Hale
Parmelia trabeculata Ahti, Acta Bot. Fenn. 70: 54 (1966).
PARATYPE. Canada: Ontario, Kenora District, Winisk. On trunks
of Populus balsamifera. T. Ahti 4669, 7 August 1958 (CANL
14265).
= Melanelia trabeculata (Ahti) Essl.
Parmelia virginica Hale, Contr. U.S. Natl. Herb. 36(4): 186
(1964).
ISOTYPE. United States: Virginia, Page Co., Hawksbill,
Shenandoah National Park. On fir trees. M.E. Hale 21592, 26.
June 1962 (CANL 31054).
= Hypotrachyna virginica (Hale) Hale
Parmeliella concinna Lamb, Farlowia 4: 431 (1955).
HOLOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Verde near
Futalaufquén. On a carbonised trunk in open forest. I.M. Lamb
5879, 1 November 1950 (CANL 2629).
Parmeliella granulata Lamb, Farlowia 4: 429 (1955).
HOLOTYPE. Argentina: Prov. Neuquén, Parque Nacional Lanén,
Lago Quillen. O. Kuhnemann 791 p.p., 10 November 1943 (CANL
2841).
Parmotrema demethylmicrophyllinicum Elix, Lich. Austral. Exs.
PASC... 9,NO.a.14 . (1986 )..
ISOTYPE. Australia: Queensland, Haning Pinnacle (Dempseys
Peak), 7 km E of Mt. Garnet. On granite rocks. J.A. Elix & H.
Streimann 16844, 29 June 1984 (CANL 95337). Elix: Lich.
Austral. Exs. No. 114.
Parmotrema queenslandense Elix, Lich. Austral. Exs. Fasc. 5,
No. 116 (1986).
ISOTYPE. Australia: Queensland, near the summit of Mt.
Leswell, 32 km S of Cooktown. On granite rocks. J.A. Elix &
He Streimann 17365, 5 July, 1984..(CANL 95339) .. Elax: Lich,
Austral. Exs. No. 116.
Parmotrema indicum Hale, Mycotaxon 5: 436 (1977).
ISOTYPE. India: Tamil Nadu, Silver Cascade, Kodaikanal, Palni
Hills. Ad saxa in vicinitate fluminis. M.E. Hale 43874,
January 1975 (CANL 69492). Vézda: Lich. Sel. Exs. No. 1573.
Peltigera kristinssonii Vitik., Ann. Bot. Fenn. 22: 291
(1985).
PARATYPE. United States: Colorado, Boulder Co., Boulder
Canyon. W.A. Weber, 23 September 1975 (CANL 58199). Weber:
332
Lich. -Exs./Colo. Now 484.
Peltigera malacea f. hasimotoi Asah. & Inum., Acta Phytotax.
Geobot. 12: 8 (1943).
ISOTYPE. Japan: Prov. Etchu, Mt. Kaminotake. Over mosses.
Hasimoto & Asahina, 24 July 1936 (CANL 20052). Kurokawa:
Lich. Rar. Crit. Exs: No. 43%.
Peltigera pacifica Vitik., Ann. Bot. Fenn. 22: 294 (1985).
ISOTYPE. Canada: British Columbia, 10 km N of Kitsumkalum
Lake on E branch road. Terricolous. T. Goward 81-2025, 24
August 1981 (CANL 104553).
Peltula cylindrica Wetm., Ann. Missouri Bot. Gard. 57: 182
(1970).
ISOTYPE. United States: Georgia, Dekalb Co., Mt. Arabia, 20
km E von Atlanta. C.M. Wetmore 19264, 1969 (CANL 102538).
Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 33.
Pertusaria gymnospora Kantvilas, Lichenologist 22: 296
(1990).
PARATYPE. Australia: Tasmania, forest around Lake Judd, SW
Tasmania. On dead wood, soil, rotten leaves. G. Bratt 73/900,
28 July 1973 (CANL 58194). Weber: Lich. Exs. Colo. No. 479.
Pertusaria neoscotica Lamb, Rep. (Annual) Natl. Mus. Canada,
1952-537, Bulls 1323"284')5(1954)%
HOLOTYPE. Canada: Nova Scotia, Cape Breton Island, Baddeck.
On trunk of trees. J. Macoun 1953, p.p. 9 July 1898 (CANL
11479).
Pertusaria paramerae A. Crespo & Vézda, Anales Jord. Bot.
Madrid 41: 252, 1984 (1985).
ISOTYPE. Spain: Prov. Soria, prope vicum judes, Paramera. Ad
corticem Juniperi thuriferae. A. Crespo & A. Vézda, 24 May
1983 (CANL 89185). Vézda: Lich. Sel. Exs, No. 1979.
Pertusaria santamonicae Dibben, Publ. Biol. Geol. Milwaukee
Public Mus. 5: 67 (1980).
PARATYPE. United States: California, San Mateo Co., San
Francisco, below Spillway of San Andres Lake. W. Jordan 769,
13 September 1967 (CANL 27071).
Pertusaria suboculata Brodo & Dibben, Publ. Biol. Geol.
Milwaukee Public Mus. 5: 72 (1980).
HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands,
Moresby Island, Tasu. On Alnus sinusta. I.M. Brodo 12825, 28
July 1967 (CANL 30938).
PARATYPES. Canada: moresby Island. I.M. Brodo 10896, 10980 &
14210 (CANL 29029, 27485 & 36493); United States: Oregon.
C.E. Fellows 32934 (CANL 38171).
Pertusaria subpupillaris Vézda, Vézda: Lich. Sel. Exs. Fasc.
20: 5 (1966).
ISOTYPE. United States: California, Mendocino Co., Caspar. On
branches of Pinus muricata. H. Sharsmith & I. Tavares 725, 25
333
July 1961 (CANL 36511). Vézda: Lich. Sel. Exs. No. 491.
= P. glaucomela (Tuck.) Nyl.
Pertusaria subvelata G. kK. Merrill, Bryologist 11: 111
(1908).
ISOTYPE. United States: Alaska, Skagway. On birch bark. J.
Macoun 87, 4 September 1902 (CANL 11603).
= P. panyrga (Ach.) Massal.
Pertusaria sulcata Dibben, Publ. Biol. Mulwaukee Public Mus.
62073) (1980)-
ISOTYPES. United States: Maine, North Lubec. On rail fences.
C.E. Cummings & E.A. Teller, 3 July 1893 (CANL 19548 &
68955). Cummings: Dec. N. Amer. Lich. No. 136 & Lich.
Boreali-Amer. No. 55.
Pertusaria victoriana Lamb, Farlowia 4: 463 (1955).
HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel
Huapi, Isla Victoria. On a large rock. I.M. Lamb 5817, 26
January 1950 (CANL 10590).
Phaeocalicium asciiforme Tibell, Symb. Bot. Upsal. 27(1): 200
(1987).
ISOTYPE. New Zealand: South Island, Westland, Arthur's Pass
National Park, 3 km NNE of Otira. On branches of Coprosma sp.
L. Tibell 9944b (CANL 99801). Tibell: Caliciales Exs. No.
168.
Phaeopeccania australiensis Henssen, Henssen: Lich. Cyanoph.
Fungi Sax. Exs. Fasc. 2: 10 (1990).
ISOTYPE. Australia: Kiistenebene zwischen Denham und Nanga; in
Vertiefungen auf horizontalen Felsplatten aus Muschelkalk. A.
Henssen 31366a, 31 August 1987 (CANL 102556). Henssen: Lich.
Cyanoph. Fungi Sax. Exs. No. 49.
Phloeopeccania hispanica Henssen & Willems, Henssen: Lich.
Cyanoph. Fungi Sax. Exs. Fasc. 2: 10 (1990).
ISOTYPE. Spain: Andalusien, Sierra de Caho de Gata, Las
Negras, Nordhang am Meeresufer. A. Henssen & G. Willems
33008a, 2-4 January 1989 (CANL 102555). Henssen: Lich.
Cyanoph. Fungi Sax. Exs. No. 48.
Phaeophyscia spinellosa Kashiw., Bull. Natl. Sci. Mus. Tokyo.
Ser. B, 10: 46 (1984).
ISOTYPE. Japan: Honshu, Prov. Suruga, Fujieda-city. On rocks.
K. Sugiyama, 10 August 1974 (CANL 87342). Kurokawa: Lich.
Rar. “Cri. xs. No. 585.
Phlyctis pulveracea Lamb, Lilloa 26: 425 (1953),
HOLOTYPE. Uruguay: Dept. Florida, La Palma. On bark. W.G.
Herter 99430, 1938 (CANL 12933).
Physcia duplicorticata W. Weber & Thomson, Mycotaxon 3: 102
(1975).
ISOTYPE. United States: California, Marin Co., 1.6 km E of
junction Nicasio-Point Reyes road E of Inverness. On trunk
334
and branches of Umbellularia californica. W.A. Weber & G.
Kunkel, 12 March 1975 (CANL 58191). Weber: Lich. Exs. Colo.
No. 476.
Physcia hirsuta var. echinella Poelt, Mitt. Bot. Staatssamml.
Muinchen 4: 298 (1962).
ISOTYPE. Italy: Appennino Ligure, Liguria, A un muro un poco
ombroso, sopra Fabiano presso La Spezia. C. & J. Poelt, April
1962 (CANL 52894). Poelt: Lich. Alpium No 179.
= Phaeophyscia cernohorskyi (Nadv.) Essl.
Pilophorus cereolus var. hallii Tuck., Syn. North Amer. Lich.
ies) (ake i
ISOTYPE. United States: Oregon. E. Hall, 1871 (CANL 68789).
Reliqy Tuck .4No. (1275
= P. clavatus Th. Fr.
Placodium coralloides Tuck., Proc. Amer. Acad. Arts 6: 287
(1866).
ISOTYPE? United States: California. On coastal rocks near San
Francisco. H.N. Bolander, 1865 (CANL 68790). Relig. Tuck. No.
L2or
= Caloplaca coralloides (Tuck.) Hult.
Placopsis alphoplacoides Lamb, Lilloa 13: 234 (1947).
ISOTYPE. New Zealand: South Island, Western Bot. Distr.,
Otira Gorge, rata belt. On earth. G.E. & G.Du Rietz 2667, 15
April 1927 (CANL 12558).
= P. trachyderma (Krempelh.) P. James
Placopsis alphoplacoides var. clavifera Lamb, Lilloa 13: 236
(1947).
ISOTYPE. New Zealand: South Island, Western Bot. Distr.,
Otira Gorge. On earth. G.E. & G.Du Rietz 2668, 15 April 1927
(CANL 12630).
= P. trachyderma (Kremp.) P. James var. clavifera (Lamb) P.
James in Mark
Placopsis contortuplicata f. fuegiensis Lamb, Lilloa 13: 276
(1947).
ISOTYPE. Argentina: Tierra del Fuego, Sierra Alvear, ca. 20
km ENE of Ushuaia. On rocks in the alpine region. R.
Santesson 901, 7 November 1940 (CANL 12560).
Placopsis dusenii Lamb, Lilloa 13: 215 (1947).
ISOTYPE. Chile: Magallanes, Isla Desolaci6én, Puerta Angosto.
P. Dusén 199 p.p., 1896 (CANL 12568).
Placopsis gelidoides Du Rietz ex Lamb, Lilloa 13: 210 (1947).
ISOTYPE. New Zealand: South Island, Cass, Mt. Misery, in
upper subalpine belt. G.E. & G.Du Rietz 1468:15, 9 January
1927 (CANL 12608).
Placopsis lateritioides Lamb, Lilloa 13: 222 (1947).
ISOTYPE. New Zealand: South Island, Cass, Mt. Misery, in
upper subalpine belt. On rocks. G.E. & G.Du Rietz 1468:16, 9
335
January 1927 (CANL 12609).
Placopsis perrugosa f. activa Lamb, Lilloa 13: 272 (1947).
ISOTYPES. Chile: Magallanes, Isla Riesco, Mina Elena. On a
stone block. R. Santesson 2023a, 29 April 1940 (CANL 12614 &
12621).
Placopsis salazina Lamb, Lilloa 13: 259 (1947).
ISOTYPE. New Zealand: Southland, Mt. Barber, above Deep Cove.
On boulder. G.E. & G.Du Rietz 2063:2, 3 March 1927 (CANL
12635).
Plagiocarpa septemseptata R. Harris, Michigan Bot. 12: 37
CL972)e
ISOTYPE. United States: North Carolina, Transylvania Co.,
Pisgah National Forest. On Fraxinus. R.C. Harris 3373, 29 May
1967 (CANL 37270).
Polyblastia cucurbitula Thomson, Bryologist 91: 89 (1988).
PARATYPES. Canada: British Columbia. Brodo 21737 & 22417
(CANL 87604 & 86571).
Polyblastia sbarbaronis Servit, Ann. Mus. Civico Storia Nat.
Giacomo Doria 66: 241 (1953).
ISOTYPE. Italy: Toscana, Vallombrosa, Saltino. Sbarbaro, July
1937 (CANL 1003).
Polyblastiopsis quercicola Brodo, Bull. New York State Mus.
mSCioeSury. 410: 147 (1968)*.
ISOTYPE. United States: New York, Suffolk Co., Shorehen,
Saint Joseph's Villa. N. Country Road. On Quercus alba. I.M.
BEOdOVZ65 19d OuULy 1961 (CANE! 2277).
= Julella fallaciosa (Stizenb. ex Arnold) R. Harris
Polycauliona coralligera Hue, Expédit. Antarct. France 1903-
05. p.10 (1908).
ISOTYPE. Antarctica: Graham Land (Palmer Peninsula), Booth
(Wandel) Island. Expédit. Antarct. France 1903-05, No. 277,
299. 1904 (CANL 17955).
= Xanthoria candelaria (L.) Th. Fr.
Porina mangiferae Vainio, Univ. Calif. Publ. Bot. 12(1): 14
(1924).
ISOTYPE. Tahiti: Society Islands, Fautaua Valley. On leaves
of Mangifera. W.A. Setchell and H.E. Parks 5091, p.p., 3 June
1922 (CANL 103926).
= P. corruscans (Rehm) R. Sant.
Porocyphus kalbarrensis Henssen, Henssen: Lich. Cyanoph.
Fungi Sax. Exs. Fasc. 2: 9 (1990).
ISOTYPE. Australia: Kalbarri National Park, Red Bluff.
Sandstein. A. Henssen 31358a, 29 August 1987 (CANL 102552).
Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 46.
Porpidia carlottiana Gowan, Bryologist 92: 39 (1989).
HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands,
336
Graham Island, Port Lewis. On rock. I.M. Brodo 10489, 26 June
1967 (CANL 27433).
Porpidia thomsonii Gowan, Bryologist 92: 54 (1989).
ISOTYPE. Canada: Northwest Territories, District of Keewatin,
North end of Lake Ennadai. On boulders. J.W. Thomson 20416,
22 July 1960 (CANL 78044).
Protoparmelia loricata Poelt & Vézda, Vézda: Lich. Sel. Exs.
Fasc. 92: .5 (1989);
ISOTYPE. Austria: Tirolia orientalis, alpes dicti Hohe
Tauern, in valle Umbaltai. J. Poelt & R. Tiirk, 29 August 1988
(CANL 99233). Vézda: Lich. Sel. Exs. No. 2292.
Pseudocyphellaria anomala Brodo & Ahti, Mycotaxon 28: 95
(1987).
HOLOTYPE. United States: Washington. On trees. A.S. Foster,
April 1908 (CANL 20272).
Pseudocyphellaria exanthematica Lamb, Farlowia 4: 435 (1955).
HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lago Frias.
On mossy trunk of Coihue. I.M. Lamb 5986, 11 November 1950
(CANL 3313).
Pseudocyphellaria hirsuta f. leucosticta Lamb, Farlowia 4:
436 (1955).
ISOTYPE. Argentina: Patagonia, Prov. Chubut, Lago
Futalaufquen. O. Ktihnemann 5449, 1941 (CANL 3334).
Pseudocyphellaria rainierensis Imsh., Mycologia 42: 749
(1950).
ISOTYPE. United States: Washington, Mt. Rainier National
Park, Ohanapecosh River above Panther Creek. H.A. Imshaug
598, 23 July 1948 (CANL 3369).
Pseudoparmelia crystallicola Kalb & Hale, Kalb: Lich. Neotr.
Fasc., 7: 14 (1983).
ISOTYPE. Brazil: Minas Gerais, Serra do Espinhacgo. Bei
Diamantina, in einer offenen Felssteppenlandschaft. K. Kalb
& G. Plébst, 17 July 1978 (CANL 85932). Kalb: Lich. Neotr.
No. 294.
Psilolechia leprosa Coppins & Purvis, Lichenologist 19: 35
(1987).
PARATYPE. Sweden: Harjedalen Prov., Tanndas par., Mt
Gruvvalen, 0.1 km NE of Lake Glimsj6n. On the walls and roof
of a cave in the old copper mine. R. Santesson 31128, 10
August 1984 (CANL 95091). Moberg: Lich. Sel. Exs. Ups. No.
16.
Psora cerebriformis W. Weber, Mycotaxon 13: 104 (1981).
ISOTYPE. United States: Colorado, Montrose Co., 4 mi. E of
Bedrock, on gypsum knolls, floor of Paradox Valley. S.
Shushan, R. Anderson & W.A. Weber 25501, 30 May 1960 (CANL
20814). Weber: Lich, Exs. Colo. No. 24.
337
Psora montana Timdal, Bryologist 89: 266 (1986).
ISOTYPE. United States: Colorado, Grand Co., Tonahutu Creek
Trail on NE end of Big Meadows. R.A. Anderson 2386, 6 July
1962 (CANL 124)
Psora pacifica Timdal, Bryologist 89: 268 (1986).
ISOTYPE. United States: California, Santa Barbara Co.,
Channel Islands, Santa Cruz Island, upper end of Islay
Canyon. On bare earth. W.A. Weber & C. Bratt, 8 January 1986
(CANL 97963). Weber: Lich. Exs. Colo. No. 664.
Psora texana W. Weber, Mycotaxon 6: 178 (1977).
ISOTYPE. United States: Texas, Kerr Co., Guadalupe River
Canyon, Hwy. 39, 22.4 km SW of Ingram, on road to Leakay. On
vertical faces of limestone cliffs. W.A. Weber, 29 April 1974
(CANL 52198).
= Xanthopsorella texana (W. Weber) Kalb & Hafellner
Psoroma internectens Lamb, Farlowia 4: 427 (1955).
ISOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Menendez.
On soil among mosses. O. Kiihnemann 4747, 1940 (CANL 3049).
Psorotichia rimosa Henssen, Henssen: Lich. Cyanoph. Fungi
Saxe SEXS -erasc. 2221 (19902
ISOTYPE. France: Languedoc, Ariége, Montagnes du Plantaurel,
anstehender Kalk an der Strasse zwischen Mas d'Azile und
Sabarat. A. Henssen 29172a, 5 June 1983 (CANL 102557).
Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 50.
Pterygiopsis australiensis Henssen, Henssen: Lich. Cyanoph.
Fungi Sax. Exs. Fasc. 2: 9 (1990).
ISOTYPE. Australia: Kalbarri National Park, Red Bluff. A.
Henssen 31357, 29 August 1987 (CANL 102550). Henssen: Lich.
Cyanoph. Fungi Sax. Exs. No. 44.
Pterygiopsis canariensis Henssen, Henssen: Lich. Cyanoph.
Fungi Sax. Exs. Fasc. 2: 8 (1990).
PARATYPE. Canary Islands: Fuerteventura,
Sukkulentenhalbwiiste, auf basisch-kristallinem Gestein. A.
Henssen 32050, 13-14 February 1988 (CANL 102548). Henssen:
Lich. Cyanoph. Fungi Sax. Exs. No. 42.
Pterygiopsis convoluta Henssen, Henssen: Lich. Cyanoph. Fungi
Sax. -Exs. Fasc. 2: 8 (1990).
ISOTYPE. Australia: Northampton Gorge, Konglomerat aus
quarzitischem Sandstein im oberen Teil der Schlucht auf
Sickerwasserflachen. A. Henssen 31373a, 1 September 1987
(CANL 102549). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No.
43.
Pterygium petersii Nyl., Syn. Meth. Lich. 1: 93 (1858).
ISOTYPE. United States: Alabama, Moulton. 7T.M. Peters, 1853
(CANL 68740). Reliq. Tuck. No. 61.
= Placynthium petersii (Tuck.) Burnham
Pyrenastrum gemmeum Tuck., Amer. J. Sci Arts Ser. 2, 25: 429
338
(1858).
SYNTYPE. United States: South Carolina, Santee Canal. H.W.
Ravenel, 1853 (CANL 68791). Reliq. Tuck. No. 129.
= Parmentaria astroidea Fée
Pyrenopsis sojakii Vézda, Folia Geobot. Phytotax., Praha 14:
205 (1979).
ISOTYPE. Iran: "Persia australis", Bandar Lengeh. Ad lapides
Calcareos locis apricis. J. Sojak, 25 April 1977 (CANL
TIE17)e
Pyrenula cinerella var. quadriloculata Fink, Minnesota Bot.
Stud w' 2257217 6atG2899)%
ISOTYPE. United States: Minnesota, Grand Portage Island. B.
Fink 85, 24 June 1897 (CANL 1384).
= ‘unknown sp. (fide R- C. Harris, "Michigan “BOC. 7 1227949
(19°73)
Pyrenula leucoplaca var. pluriloculata Fink, Minnesota Bot.
Stud. 2: 709 (1902).
SYNTYPE. United States: Minnesota, Thief River Falls. On
trees. B. Fink 829, 19 July 1900 (CANL 1335).
= Eopyrenula intermedia Coppins in Aptroot (1991).
Pyrenula macounii R. Harris, Michigan Bot. 12: 45 (1973).
HOLOTYPE. Canada: Ontario, Belleville. On trees. J. Macoun,
April 1874 ‘(CANL 1350).
Pyrenula neglecta R. Harris, Michigan Bot. 12: 45 (1973).
ISOTYPE. United States: New Hampshire, Plymouth. On beech
trees. C.E. Cummings, 26 August 1893 (CANL 19561). Cummings:
Dec. N. Amer. Lich. No. 150.
= Pyrenula pseudobufonia (Rehm) R. Harris
Pyrenula neglecta subsp. occidentalis R. Harris, Michigan
Bots §12:°'51°(1973)5
ISOTYPE. United States: Washington, Gate. On vine maple. A.S.
Foster, 30 March 1912 (CANL 20502). Merrill: Lich. Exs. No.
5 Pale
= P. occidentalis (R. Harris) R. Harris
Pyrenula nitidella var. maculata R. Harris, Weber: Lich. Exs.
Colo. Fasc. +120 eNow'4437(1975)..
ISOTYPE. United States: Texas, McLennan Co., Canyon of the
Middle Bosque River SW of Valley Mills and W of Waco. On
saplings of a smooth-barked tree. W.A. Weber, 27 May 1974
(CANL 52193). Weber: Lich. Exs. Colo. No. 443.
= P. maculata (R. Harris) R. Harris
Pyrenula shirabeicola Kurok. & S. Nakan., Mem. Natl. Sci.
Mus. TOKYO?4 : +67 VLSI
ISOTYPE. Japan: Honshu, Prov. Hida, Mt. Ontake. On trunk of
Abies veitchii. S. Kurokawa 64172, 13 August 1964 (CANL
45336). Kurokawa: Lich. Rar. Crit. Exs. No. 188.
Pyrenula thelomorpha Tuck., Gen. Lich. p. 275 (1872).
339
ISOTYPE. United States: South Carolina, Santee Canal. On Ilex
decidua. H.W. Ravenel 468 (CANL 68797). Reliq. Tuck. No. 135.
= Anthracothecium thelomorphum (Tuck.) Zahlbr.
Pyxine daedalea Krog & R. Sant., Publ. Herb. Univ. Uppsala,
Thunbergia 2: 7 (1986).
ISOTYPE. Costa Rica: Cartago Prov., 13 km SE of Cartago, 2.5
km SE of Orosi near the bridge over Rio Grande de Orosi. On
a large boulder. H. Krog & R. Santesson 29074, 10 January
1979 (CANL 95093). Moberg: Lich. Sel. Exs. Ups. No. 17.
Ramalina americana Hale, Bryologist 81: 599 (1978).
ISOTYPES. United States: New Hampshire, Plymouth. On yellow
birch. C.E. Cummings, 28 May 1891 (CANL 19451, 68913).
Cummings: Dec. N. Amer. Lich. No. 43 & Lich. Boreali-Amer.
No. 5.
Ramalina atlantica Culb., Brittonia 19: 350 (1967).
ISOTYPE. Scotland: Argyllshire, Ardnamurchan Peninsula,
Portaick. On immense boulder. W.L. Culberson 12254, July 1964
(CANL 22578).
Ramalina bicolor Mull. Arg., Flora 60: 476 (1877).
ISOTYPE. Brazil: Bahia, Blanchet (CANL 42339).
Ramalina cactacearum Follm., Nova Hedwigia 14: 256 (1967).
ISOTYPE. Chile: Prov. Antofagasta, zerstreut im Dornenwerk
von Sdaulenkakteen im Tornabenietum intricatae. I.A. Follmann-
Schrag & G. Follmann, September 1965 (CANL 33001). Follmann:
Pach mExs: Sell. No. #51.
Ramalina euxini Vézda, Folia Geobot. Phytotax., Praha 14: 205
(1979).
ISOTYPE. Bulgaria: Distr. Burgas, in litore Euxini Ponti, 5
km ad meridiem versus ab oppido Sozopol. Ad scopulos
andesiticos. A, Vézda, 20 September 1979 (1978 in protologue)
(CANL 73831). Vézda: Lich. Sel. Exs. No. 1668.
Ramalina sideriza Zahlbr., Ann. K. K. Naturhist. Hofmus. 25:
246' (1911).
ISOSYNTYPE. United States: Hawaii, Island of Kauai. On trunk
of trees. A.A. Heller, 28 June 1895 (CANL 20370). Merrill:
PLC uxs. NO-1e120.
Ramalina superfraxinea Follm. & Sanch.-Pinto, Philippia 4:
192 (1980).
ISOTYPE. Canary Islands: Tenerife, Vogelnestartig im
halbschattigen Unterkronenteil solitdrer Nadelbadume auf
jungen Lavastrémen im Ramalinetum subgeniculatae. G. Follmann
& L. Sanchez-Pinto, April 1978 (CANL 75275). Follmann: Lich.
EXS* wel No: 314"
Ramalina tenuis Fries & Tuck. in Tuck., Amer. J. Sci. Arts
Serie 2)) 2520-423 4(1858)™
ISOLECTOTYPE. United States: Texas, Mexican Bountary.
Thickets of the Blanco. C. Wright, 1849 (CANL 68759). Reliq.
340
Tuck. No. 84.
LECTOPARATYPE. United States: South Carolina, Santee Canal,
H.W. Ravenel, 1869 (CANL 68760). Reliq. Tuck. No. 85.
Ramalina tigrina Follm., Willdenowia 4: 227 (1966).
ISOTYPE. Chile: Prov. Antofagasta, gesellig auf
Erosionsrippen. I.A. Follmann-Schrag & G. Follmann, September
1965 (CANL 32975). Follmann: Lich. Exs. Sel. No. 10.
Ramonia intermedia Kalb, Kalb: Lich. Neotr. Fasc. 6: 14
(1983).
ISOTYPE. Brazil: Minas Gerais, Serra da Mantiqueira, Oberhalb
von Vila Monte Verde, etwa 30 km 6stlich von Camanducaia. In
einem Gebirgsregenwald. K. Kalb & H. Sipman, 2 July 1979
(CANL 84791). Kalb: Lich. Neotr. No. 242.
Ramonia kandlerii Kalb, Kalb: Lich. Neotr. Fasc. 6: 14
(1983). |
ISOTYPE. Brazil: Mato Grosso do Sul. Zwischen Rio Verde do
Mato Grosso und Coxim. In einem dichten Cerradado in einer
Bachschlucht. K. Kalb, 28 June 1980 (CANL 84792). Kalb: Lich.
Neotr. No. 243.
Rhizocarpon compositum Lamb, Lilloa 14: 222 (1948).
ISOTYPE. Argentina: Prov. Tucuman, Quebrada de Lules. On rock
outcrop. I.M. Lamb 5132, 5 October 1947 (CANL 18207).
= Buellia excellens H. Magn.
Rhizocarpon alaxensis Thomson, Nova Hedwigia 14: 471 (1967).
ISOTYPE. United States: Alaska, Franklin Bluffs on the
Sagavanirktok River. On rocks. J.W. Thomson, S. Shushan & J.
Koranda 10859, 30 July 1958 (CANL 44991).
Rhizocarpon atlanticum Lamb, J. Bot. 78: 132 (1940).
PARATYPE. England: Cornwall, Rocky Valley near Boscastle.
I.M. Lamb 877, 1939 (CANL 6111).
= R. constrictum Malme
Rhizocarpon barilochense Rds&nen, Arch. Soc. Zool.-Bot. Fenn.
"Vanamo" 6(2): 85 (1952).
ISOTYPE. Argentina: Patagonia, Prov. Rio Negro, Cerro
Catedral near Bariloche. On rocks. I.M. Lamb 5958, 10
February 1950 (CANL 27348).
Rhizocarpon cumulatum Thomson, Nova Hedwigia 14: 461 (1967).
ISOTYPE. United States: Alaska, Pitmegea River, 15 miles
upstream from Cape Sabine. On shaded layers at top of bluff.
J.W. Thomson 10571, 12 July 1958 (CANL 45003).
Rhizocarpon hensseniae Brodo, Biblioth. Lichenol. 38: 32
(1990).
HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands,
Graham Island, Seal Inlet in Rennell's Sound. On exposed
stone. I.M. Brodo 10264, 23 June 1967 (CANL 98871).
Rhizocarpon inimicum Poelt & Vézda, Herzogia 6: 471 (1984).
341
ISOTYPE. Spain: Catalonia, montes Montseny, Matagalls. Ad
saxa schistosa, in thallo Lecanorae rupicolae. J. Poelt & A.
Vézda, 28 May 1983 (CANL 89186). Vézda: Lich. Sel. Exs. No.
1980 (1970 in protologue).
Rhizocarpon renneri Poelt, Planta 51: 306 (1958).
ISOTYPE. Austria: Samnaungruppe, Tirol. Steilflache des
grossen Blockes am K6lner Haus. J. Poelt, July 1958 (CANL
52817). Poelt: Lich. Alpium No. 100.
Ricasolia beckettii Stirton, Trans. & Proc. New Zealand Inst.
3.075 °392°°(1897)), 2
ISOTYPE. New Zealand: Mount Bossu, Banks Peninsula. On
Podocarpus totara. T.W.N. Beckett L. 3, 1893 (CANL 3387).
= Pseudocyphellaria coriacea (T. D. Hook. & Taylor) D.
Galloway & P. James ?
Rimularia fuscosora Muhr & T@nsberg, Nord. J. Bot. 8: 649
(1989).
ISOTYPE. Sweden: Vdsterbotten Prov., Skelleftea par., ca. 3
km NE of Mt Stora Blabergsliden, by the small stream
Djupgravbacken. On Alnus incana. L.-E. Muhr 7540, 8 August
1984 (CANL 102833). Moberg: Lich. Sel. Exs. Ups. No 94.
Rinodina afghanica M. Steiner & Poelt, Pl. Syst. Evol. 155:
139 (1987).
ISOTYPE. Afghanistan: Prov. Samangan, 1 km W des Passes
Kotal-Mirza Atbili. M. Steiner Ste 50/3, 6 June 1970 (CANL
104080). Steiner & Poelt: Lich. Afghanica No. 46.
Rinodina ascociscana Tuck., Gen. Lich. p. 124 (1872).
ISOSYNTYPE. United States: New Hampshire, White Mountains. E.
Tuckerman (CANL 23267). Reliq. Tuck. No. 86.
Rinodina calculiformis W. Weber, Bryologist 67: 473 (1964).
ISOTYPE. Mexico: Baja California, Isla Guadalupe, Melpomene
Cove. On volcanic scoria. W.A. Weber & C.J. McCoy, 21 April
1963 (CANL 20883). Weber: Lich. Exs. Colo. No. 94.
Rinodina dakotensis H. Magn., Bot. Not. 1947: 40 (1947).
ISOTYPES. United States: South Dakota, Brookings. On plum
trees. 1T.A. Williams, 11 May 1895 (CANL 19580, 69031).
Cummings: Dec. N. Amer. Lich. No. 169b & Lich. Boreali- Amer.
No. 163b.
Rinodina humilis H. Magn., Bot. Not. 1939: 132 (1939).
ISOTYPE. Sweden: Vastergdétland, Bjdérketorp, Stenbacka. On
moist stones. A.H. Magnusson, 26 March 1934 (CANL 20147).
Magnusson: Lich. Sel. Scand. Exs. No. 324.
Rinodina milliaria f. obscura H. Magn., Bot. Not. 1947: 19
(1947).
ISOTYPE. United States: Maine, Rockland. On bark of young
trees. G.K. Merrill 163, 21 April 1910 (CANL 20406). Merrill:
Lich. .Exs< No. 163)
= R. dakotensis H. Magn.
342
Rinodina oregana H. Magn., Bot. Not. 1947: 49 (1947).
ISOTYPES. United States: Oregon, Forest Grove. On Acer
macrophyllum. A.R. Sweetser, December 1899 (CANL 19739,
69098). Cummings: Dec. N. Amer. Lich. No. 330 & Lich.
Boreali-Amer. No. 262.
Rinodina pallida H. Magn., Bot. Not. 1939: 313 (1939).
ISOTYPE. Sweden: Vdstergétland, Broddetorp, Fjdllakra. On a
bridge of sandstone. A.H. Magnusson, 10 June 1938 (CANL
20148). Magnusson: Lich. Sel. Scand. Exs. No. 325.
Rinodina sabulosa Tuck., Lich. Calif. p. 21 (1866).
ISOTYPE. United States: California. H.N. Bolander, 1865 (CANL
68761) Reliq.w2uck. eNO. S7:
= R. conradii Kdérber
Roccella galapagoensis Follm., Nova Hedwigia 15: 337 (1969).
ISOTYPE. Ecuador: Galapagos Islands, Isla Santa Cruz. On
seaward vertical sides of bouders. W.A. Weber, 13 February
1964 (CANL 20901) Weber: Lich. Exs. Colo. No. 112.
Roccellina luteola Follm., Nova Hedwigia 14: 239 (1967).
ISOTYPE. Chile: Prov. Antofagasta, koloniebildend an
Stirnfldchen und Uberhdngen von Diorithéckern im Roccelletum
portentosae. G. Follmann, September 1965 (CANL 33004).
Follmann: Lich. Exs. Sel. No. 54.
Roccellina olivacea Follm., Philippia 4: 111 (1979).
ISOTYPE. Chile: Prov. Coquimbo, gesellig an schattseitigen,
durch Gischtstaub beeinflussten Stirnfldchen und Uberhdngen
von Dioritfelsen im Roccelletum portentosae. Ktistenformation
nérdlich der Limarimtindung. G. Follmann, September 1965 (CANL
71806). Follmann: Lich. Exs. Sel. No. 296.
Sagenidiopsis merrotsii R. W. Rogers & Hafellner,
Lichenologist 19: 402 (1987).
ISOTYPE. Australia: New South Wales, Border Ranges National
Park, Brindle Creek. Ad truncum Nothofagi moorei. J.
Hafellner, P. Merrotsy & R. Rogers, 30 August 1986 (CANL
97328). Vézda: Lich. Sel. Exs. No. 2226.
Sarrameana tasmanica Vézda & Kantv., Lichenologist 20: 179
(1988).
ISOTYPE. Australia: Tasmania, Little Fisher River. Ad
corticem arboris. G. Kantvilas & J. Jarman 63/87, 15 May 1987
(CANL 97302). Vézda: Lich. Sel. Exs. No. 2200.
Schistophoron variabile Tibell, Lichenologist 14: 242 (1982).
ISOTYPE. Costa Rica: Lim6én, 1 km NW of Pto. Lim6én. On trunk
of Cocos nucifera. L. Tibell 8514, 9 January 1979 (CANL
87469). Tibell: Caliciales Exs. No. 96.
Schizopelte californica Th. Fr., Flora 58: 143 (1875).
ISOTYPE. United States: California. G. Fisen, 1874 (CANL
2122).
= Combea californica (Th. Fr.) Follm. & Geyer
343
Setaria trichodes Michaux, Fl. Boreal.-Amer. 2: 331 (1803).
ISOTYPE. Canada: Ad ramulos putridos (CANL 34981).
= Bryoria trichodes (Michaux) Brodo & D. Hawksw. subsp.
trichodes.
Skyttea tephromelarum Kalb & Hafellner, Kalb: Lich. Neotr.
Frascw 103 "15 (1988)':
ISOTYPE. Kenya: Nanyuki District zwischen Naro Moru und
Nanyuki. K. Kalb & A. Schrégl, 18 August 1985 (CANL 102737).
Kalb: Lich. Neotr. No. 449.
Sphaerophorus notatus Tibell, Publ. Herb. Univ. Uppsala 10:
9 (1982).
ISOTYPE. New Zealand: Otago, Mount Aspiring National Park, 10
km NNE of Makaroa. On trunk of Nothofagus menziesii. L.
Tibell 10604, 23 January 1981 (CANL 82051). Tibell:
Caliciales Exs. No. 73.
Sphaerophorus ramulifer Lamb, Farlowia 4: 426 (1955).
HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lago Frias.
On trunk of Fitzroya near the lake. I.M. Lamb 5977, 11
February 1950 (CANL 1751).
Squamaria configurata f. diffracta Oxner, Ukrayins'k. Bot.
Zhurns.4353) (1928)
ISOTYPE. Ukraine: Ditio Kirowogradensis. Ad safa granitica ad
flum. Subakleji in viciniis opp. Kirowograd. A. Oxner, 22 May
1926 (CANL 11909).
= Lecanora laatokkaensis (R&sdnen) Poelt
Stephanocyclos henssenianus Hertel, Hertel: Lecideaceae Exs.
Fasc. 5, No. 96 (1983).
ISOTYPE. Subantarctic Islands: Marion Island, Nellie Humps,
zwischen Junior's Kop und Meteorologischer Station. Niedrige,
bizarre Lava-Felsen im moorigen Geldnde. H. Hertel, 13 April
1982 (CANL 85425). Hertel: Lecideaceae Exs. No. 96.
Stereocaulon alpinum var. erectum Frey, Rabenh. Kryptog. Fl.
9 Abt. 4, 1. Halfte: 164 (1932).
ISOLECTOTYPES. Switzerland: Graubiinden, Engadin, Samaden,
Kies- und Sandalluvionen des Inn nahe beim Bahnhof. E. Frey
1636, 1931 (CANL 77611, 63390). Poelt: Pl. Graecenses No.
232%
Stereocaulon arbuscula var. aberrans Asah., J. Jap. Bot. 19:
282 (1943).
ISOLECTOTYPE. Japan: Honshu, Prov. Musashi, Nippara. I.
Sasaki, 1940 (CANL 48276).
= Leprocaulon arbuscula (Nyl.) Nyl. (Ch. strain II.)
Stereocaulon argodes Nyl., Compt. Rend. Hebd. Séances Acad.
SGPIES3"87 61876) ¢
ISOTYPE. New Zealand: Campbell Island. Filhol, 1874 (CANL
67862)"
= S. argus J. D. Hook. & Taylor
344
Stereocaulon argus var. stenospermum Lamb, Farlowia 4: 460
(1955).
HOLOTYPE. Argentina: Prov. Rio Negro, near foot of Frias
Glacier S. of Lago Frias. On dry exposed rocks. I.M. Lamb
6088, 1950 (CANL 63410).
ISOTYPE. (CANL 77621). Poelt: Pl. Graecenses No. 242.
= S. stenospermum (Lamb) Lamb
Stereocaulon azulense Yoshimura & W. Weber, Mycotaxon 27: 494
(1986).
ISOTYPE. Ecuador: Galapagos Islands, Isla Isabela, SW coast,
Cerro Azul. On boulders. W.A. Weber & H. Beck, 17 January
1984 (CANL 97944). Weber: Lich. Exs. Colo. No. 645.
Stereocaulon botryosum f. depressum Frey, Rabenhorsts
Kryptog "Flos 9 (4) Di 26 1Clo3 2).
ISOLECTOTYPE. Norway: Nordland, Umbukten. On sunny places.
A.H. Magnusson 9040, 1924 (CANL 70674).
= S. depressum (Frey) Lamb
Stereocaulon botryosum f. pygmaeum H. Magn., Ark. Bot. 33A:
91 (1946).
ISOTYPE. Sweden: Lycksele Lappmark, Tarna, Brakkffallet. On
irrigated rock. A.H. Magnusson 8084, 1924 (CANL 75018).
= S. spathuliferum Vainio f. pygmaeum (H. Magn.) Lamb
Stereocaulon capense Lamb, Lilloa 26: 416 (1953).
HOLOTYPE. South Africa: Western Cape Province, Tulbagh
Division, Great Winterhoek. On rock face. E. Esterhuysen
19862 p.p., 1951 (CANL 63391).
= S. corticatulum Nyl. var. capense (Lamb) Lamb
Stereocaulon colensoi var. reagens R&dsdnen, Ann. Bot. Soc.
Z00l.=Bot.. Fenn .."Vanamot 22624 2(1932)r,
ISOSYNTYPE? Chile: Fuegia occ., Fjordo Finlandia, supra
rupium. H. Roivainen, 1929 (CANL 70923).
= S. glabrum (MUll. Arg.) Vainio
Stereocaulon condensatum var. sorediatum Harm., Bull. Acad.
Int. .Géoqr.) Bot. 22: 5157161913) '
ISOTYPE. France: Hérault, La Salvetat. Sur des rochers
schisteux, terreux. F. Marc (CANL 71016). Claud. & Harm.:
Lich. Gall. Praecip. Exs. No. 474.
Stereocaulon coralloides Fr., Fr.: Lich. Suec. Exs. No. 118
(1817).
ISOTYPE. Sweden: no locality or date designated (CANL 70733).
= S. dactylophyllum Floérke
Stereocaulon coralloides var. flabellatum Frey, Rabenhorsis
Kryptog. ‘Fl. 9 (4,1)3,196 (1932).
ISOLECTOTYPE. Germany: Bayen, Bohmerwald, Bayrisch-
Eisenskein, Hochbergplatte, schatlig. J. Hillmann, 2 October
1930 (CANL 70680)
= S. dactylophyllum Flérke var. flabellatum (Frey) Grumn.
345
Stereocaulon coralloides var. occidentale H. Magn., Géteborgs
Kungl. Vetensk. Samhdlles Handl. Ser. 4, 30: 27 (1926).
ISOLECTOTYPE. Sweden: Bohuslan, Dragsmark. A.H. Magnusson,
1915 (CANL 70696).
= S. dactylophyllum Flérke var. occidentale (H. Magn.) Grumn.
Stereocaulon cornutum var. corallizans Lamb, J. Hattori Bot.
Babe 43082 5270-0197 7)2
ISOTYPE. Costa Rica: Prov. Heredia, Cerro Central de Zurqui.
C.W. Dodge, J. Valerio R., W.S. Thomas & R. Valerio 6119,
1929 (CANL 63389).
Stereocaulon corticatulum var. complanatum Lamb, J. Hattori
Bots? Labs 4327 289¢ (1977).
ISOTYPE? New Zealand: Stidinsel, Otago, Taieri Mouth,
Kuistenfelsen. J. Murray 9742 (1421 in protologue), 1958.
(CANL 77612). Poelt: Pl. Graecenses No. 233.
Stereocaulon corticatulum var. procerum Lamb, Farlowia 4: 462
(1955).
HOLOTYPE. Argentina: Prov. Rio Negro, between Lago Frias and
Frias Glacier. On rocks. I.M. Lamb 6086, 1950 (CANL 63400).
ISOTYPE. (CANL 77619).
= S. melanopotamicum Lamb
Stereocaulon crambidiocephalum Lamb, J. Hattori Bot. Lab. 43:
290 (1977).
ISOTYPE. Peru: Dept. Cuzco, Prov. Paucartambo, Llulluchayocc.
On rock. C. Vargas 4301, 1944 (CANL 63392).
Stereocaulon dendroides Asah., J. Jap. Bot. 36: 231 (1961).
ISOTYPE. Japan: Shikoku, Mt. Kenzan. F. Fujikawa, 1934 (CANL
63393).
Stereocaulon denudatum var. umbricola ("umbricolum") Frey,
Rabenhorsts Kryptog. Fl. 9 (4,1): 134 (1933).
ISOTYPE. Switzerland: Berner Oberland, Grimsel, way towards
Nagelisgraéatli. On granite rock. E. Frey 1429, 1930 (CANL
63414).
= S. vesuvianum Pers. var. nodulosum (Wallr.) Lamb f.
umbricola (Frey) Lamb
Stereocaulon dusenii Lamb, J. Hattori Bot. Lab. 43: 292
(1977).
ISOTYPE. Chile: Magallanes, Isla Desolaci6én, Puerto Angosto.
On rocks. P. Dusén 186, 1896 (CANL 63394).
Stereocaulon esterhuysenae Lamb, Lilloa 26: 417 (1953).
HOLOTYPE. South Africa: Western Cape Province, Tulbagh
Division, Great Winterhoek. On a rock face. E. Esterhuysen
19862 p.p., 1951 (CANL 63395).
Stereocaulon evolutoides var. paschaleoides Hav., Arbok.
Univ. Bergen, Naturvitensk. Rekke 12: 1720 (1954).
ISOTYPE. Norway: Granvin, Gamladaemmo, near Skalsaete. On
gneiss boulders. J. Havaas (CANL 63409).
346
= S. saxatile H. Magn. f. paschaleoides (Hav.) Lamb
Stereocaulon exutum Nyl., Lich. Japon. p. 18 (1890).
ISOTYPE. Japan: Fusijama. E. Almquist, 1879 (Vega Exped.,
1878-1880) (CANL 70910).
Stereocaulon fastigiatum f. confluens H. Magn., Gdteborgs
Kungl. Vetensk. Samhalles Handl., Ser. 4, 30: 35 (1926).
ISOTYPE. Sweden: Torne Lappmark, par. Jukkasjarvi,
Kopparasen. A.H. Magnusson 6073, 1921 (CANL 70726).
= S. depressum (Frey) Lamb
Stereocaulon fastigiatum var. dissolutum H. Magn., Gdéteborgs
Kungl. Vetensk. Samhalles Handl. Ser. 4, 30: 36 (1926).
LECTOPARATYPE. Norway: Nordland, Umbukten, Krabbfjdall. A.H.
Magnusson 9040, 1924 (CANL 75616).
= S. spathuliferum Vainio f. congestum (H. Magn.) Lamb
Stereocaulon glareosum var. brachyphylloides Lamb, J. Wash.
ACAG siSCive 4 LOG m Loo Ljts
ISOTYPES. United States: Alaska, Brooks Range, Anaktuvuk
Pass. On soil. G.A. Llano & N. Weber 527, 1949 (CANL 63397,
77614). Poelt: Pl. Graecenses No. 235.
Stereocaulon halei Lamb, J. Hattori Bot. Lab. 43: 294 (1977).
ISOTYPE. Malaysia: Pahang, unterhalb Ganong Brinchang. M.E.
Hale, 1965 (CANL 77615). Poelt: Pl. Graecenses No. 236.
Stereocaulon intermedium f. compactum Lamb, J. Hattori Bot.
Vaberssceccol oy).
ISOTYPES. United States: Alaska, Aleutian Islands, Rat
Island, Amchitka, near Consantine Harbor. G.A. Llano 1277,
1949 (CANL 63385, 77617). Poelt: Pl. Graecenses No. 238.
Stereocaulon japonicum var. subfastigiatum Asah., J. Jap.
Botte 35:1289) (196077
ISOTYPE. Japan: Honshu, Prov. Owari, Kiso River, Inuyama. Y.
Asahina 1201, 1938 (CANL 63398).
Stereocaulon japonicum var. tokioense Lamb, J. Jap. Bot. 44:
265 (1969).
HOLOTYPE. Japan: Honshu, Tokyo. On stone wall of the
University. Y. Asahina 650, 1943 (CANL 63399).
Stereocaulon leprocauloides Lamb, Bryologist 74: 193 (1971).
ISOTYPES. Papua New Guinea: Morobe District, Mt. Kaindi near
Wau. W.A. Weber & D. McVean, 19 June 1968 (CANL 33463,
77618) .. .Weber:* ) Lichy/ -Exs.°"Colov’ =NovF “299% ~Poelt:” “Pi:
Graecenses No. 239.
Stereocaulon melanopotamicum Lamb, J. Hattori Bot. Lab. 43:
2961977 )r.
(see Stereocaulon corticatulum var. procerum Lamb).
Stereocaulon meyeri f. tucumanum Lamb, J. Hattori Bot. Lab.
As AO BT. hee
347
ISOTYPE. Argentina: Prov. Tucuman, Dept. Chicligasta, between
La Cascada and Las Cuevas. On rock. T. Meyer, 1949 (CANL
63401).
Stereocaulon montagneanum Lamb, J. Jap. Bot. 40: 272 (1965).
ISOTYPE. Malaysia: Cameron Highland, On exposed rocks. M.
Togashi 62236, 1-3 February 1962 (CANL 34168). Kurokawa:
Lich. tRar. (CritstExss) Now142:
Stereocaulon myriocarpum var. altaicum Lamb, J. Hattori Bot.
Lab. 43: 225 (1977).
ISOTYPE. Kazakhstan: Siberia, Gub. Semipalatinsk, Altai Mts.,
Katon-Karagai. P. Krylov & L. Sergievskaya 8, 1928 (CANL
74958).
Stereocaulon nanodes f. schadeanum Lamb, J. Hattori Bot. Lab.
Ape e242) (197.7) i
ISOTYPE. Germany: Sachsen, Freiberg. A. Schade 1767, 1927
(CANL 63402).
Stereocaulon novoarbuscula Asah., J. Jap. Bot. 19: 283
(1943). .
ISOLECTOTYPE. Japan: Shikoku, Prov. Tosa, Mt. Otoyama. F.
Fujikawa, 1931 (CANL 48290).
= Leprocaulon pseudoarbuscula (Asah.) Lamb & Ward
Stereocaulon octomerelloides Asah., J. Jap. Bot. 45: 68
(1970).
ISOTYPE. Japan: Honshu, Prov. Etigo, Sasagamine Pasture. Y.
Asahina 1521, 1949 (CANL 63403).
= S. octomerellum Miill. Arg.
Stereocaulon papuanum Lamb, J. Hattori Bot. Lab. 43: 225
(197%) &
ISOTYPE. Papua New Guinea: Central Division, Mt. Albert
Edward. L.J. Brass 4394a, 1933 (CANL 63404).
Stereocaulon paschale var. alpinum f. flabellans Lamb,
Farlowia 4: 457 (1955).
HOLOTYPE. Argentina: Prov. Rio Negro, Cerro Rigi near Lago
Frias. On the ground between rocks and stones. I.M. Lamb
5787, 1950 (CANL 63396).
ISOTYPE. (CANL 77613). Poelt: Pl. Graecenses No. 234.
= S. glabrum (MUtill. Arg.) Vainio f. flabellans (Lamb) Lamb
Stereocaulon paschale f. colligatum Lamb, Rep. (Annual) Natl.
Mus. Canada, 1952-53, Bull. No. 132: 273 (1954).
HOLOTYPE. Canada: Nova Scotia, Victoria Co., Cape Breton
Island, Ingonish, Middle Head. On granitic rocks. I.M. Lamb
6946, 29 July 1952 (CANL 10301).
= S. paschale (L.) Hoffm. f. paschale
Stereocaulon paschale subsp. evolutoides f. laxatum Lamb,
Rep. (Annual) Natl. Mus. Canada, 1952-53, Bull. No. 132: 275
(1954).
ISOTYPE. Sweden: Varmland, Sillerud, Jarnsj6n. A.H.
348
Magnusson, 1912 (CANL 71061).
Stereocaulon paschale v. evolutoides f. sorediatum H. Magn.,
G6teborgs Kungl. Vetensk. Samhalles Handl. Ser. 4, 30: 36
(1926).
ISOTYPE. Sweden: Vdarmland, par. Sillernd, Jarnsj6n. A.H.
Magnusson, 1912 (CANL 71078).
= S. saxatile H. Magn. f. sorediatum (H. Magn.) Lamb
Stereocaulon paschale var. serpens Th. Fr., De Stereoc. et
Pilophor. Comment p. 33 (1857).
ISOTYPE. Sweden: Uppsala, Vitulfsberg. Th. Fries, 1852 (CANL
70990).
= S. paschale (L.) Hoffm.
Stereocaulon patagonicum Lamb, Farlowia 4: 454 (1955).
HOLOTYPE. Argentina: Prov. Chubut, Lago Menéndez, W. end. On
sandy gravelly shore of lake. I.M. Lamb 5790, 1950 (CANL
63405).
ISOTYPE. (CANL 77616). Poelt: Pl. Graecenses No. 237.
= S. glabrum (MUll. Arg.) Vainio
Stereocaulon patagonicum f. subirregulare Lamb, Farlowia 4:
456 (1955).
HOLOTYPE. Argentina: Prov. Rio Negro, Lago Frias, Cerro Rigi,
near summit. In cleft between stones. I.M. Lamb 5789, 1950
(CANL 63406).
= S. glabrum (MUll. Arg.) Vainio
Stereocaulon philippinense R&sd&dnen, Arch. Soc. Zo0ol.-Bot.
Fenn. "Vanamo" 3: 79 (1949).
ISOTYPE. The Philippines: Negros, Canlaon Volcano. Terricola.
E.D. Merrill 6874, April 1910 (CANL 71010).
Stereocaulon pileatum f. macrum H. Magn., Géteborgs Kungl.
Vetensk. Samhdlles Handl. Ser. 4, 30: 70 (1926).
ISOTYPE. Sweden: Ljung, Kolbengtsered. On rock. A.H.
Magnusson 9513, 13 August 1925 (CANL 71041).
= S. pileatum Ach.
Stereocaulon pileatum var. nipponicum Lamb, J. Hattori Bot.
LAD Ina Ss ees iy
HOLOTYPE. Japan: Jiushiu, Prov. Higo, Aida. K. Maebara, 1927
(CANL 63407).
Stereocaulon proximum var. gracilius Miill. Arg., Rev. Mycol.
(Toulouse) 1: 164 (1879).
ISOTYPE? Colombia "Nova Granata": La Horqueta prope Dolores.
André (coll. no. = 2813 p.p. in holotype), 1876 (CANL 77638).
= S. ramulosum (Sw.) Rausch. var. gracilius (Miill. Arg.) Lamb
Stereocaulon pseudoarbuscula Asah., J. Jap. Bot. 19: 282
(1943).
ISOLECTOTYPE. Japan: Honshu, Prov. Musashi, Titibu. Y.
Asahina 1249, 1933 (CANL 48292).
= Leprocaulon pseudoarbuscula (Asah.) Lamb
349
Stereocaulon pseudomassartianum Lamb, Bot. Jahrb. Syst. 86:
246 (1967).
ISOSYNTYPES. The Philippines: Luzon, Mountain Prov., Ifugao
Subprov. On trail between Banaue and Mt. Polis. On earth.
A.W. Herre, 1923 (CANL 71040, 77620). Poelt: Pl. Graecenses
No. 241.
Stereocaulon pygmaeum Vainio, Voyage S. Y. Belgica, Bot. p.
15 (1903).
ISOTYPE. Antarctica: Graham Land (Palmer Peninsula), Cape
Anna Osterrieth. Exped. Antarctica Belge 1897-99, No. 201.
(CANL 12807).
= Lecania brialmontii (Vainio) Zahlbr.
Stereocaulon ramulosum var. pulvinare f. crebratum Lamb, J.
Hatcoriebots Labosd43cee85in(1977))%
ISOTYPE. New Zealand: South Island, Port Chalmers near
Dunedin. On rock. J.S. Thomson D.15, T. 696 (CANL 63408).
Stereocaulon speciosum Lamb, Farlowia 4: 458 (1955).
HOLOTYPE. Argentina: Prov. Chubut, Lago Menendez. On mossy
gravelly soil. I.M. Lamb 6069, 1950 (CANL 63387).
ISOTYPE. (CANL 77623). Poelt: Pl. Graecenses No. 244.
= §. verruciferum Nyl.
Stereocaulon speciosum var. surreptans Lamb, Farlowia 4: 459
G1955)%
HOLOTYPE. Argentina: Prov. Rio Negro, Cerro Rigi near Lago
Frias, near summit. On detritus. I.M. Lamb 6098, 1950 (CANL
63388).
ISOTYPE. (CANL 77624). Poelt: Pl. Graecenses No. 245.
= S. verruciferum Nyl. var. surreptans (Lamb) Lamb
Stereocaulon subalbicans Lamb, Bryologist 60: 220 (1957).
ISOTYPE. Chile: Prov. Coquimbo, La Serena, Cerro Los Loros.
R. Santesson 2530, 1940 (CANL 48278).
= Leprocaulon subabicans (Lamb) Lamb & Ward
Stereocaulon subcoralloides f. sorediascens Lamb, J. Hattori
Bot. Lab. 43: 235 (1977).
HOLOTYPE. Russia: Karelia ladogensis, Kurkijoki, Kuuppala,
Jaavuori. On rock. V. R&s&nen, 1931 (CANL 63411).
Stereocaulon tennesseense H. Magn., Ark. Bot. 30A: 48 (1941).
ISOTYPE. United States: Tennessee, Great Smoky Mountains,
near Alum Cave. On moist rocks. G. Degelius, 1939 (CANL
75021).
Stereocaulon tennesseense var. nigrofastigiatum Lamb, J.
MaAGlLori sSOG lab. b4oc 20 Om CLO Ia):
HOLOTYPE. United States: New York State, Adirondacks, Mt.
Colden near Lake Placid. On rocks. J.L. Lowe, 1952 (CANL
63412).
ISOTYPE. (CANL 77622). Poelt: Pl. Graecenses No. 243.
Stereocaulon tomentosum var. compactum Frey, Rabenhorsts
350
Krypts RL. -9°7°(471)2"187,"(1932)".
ISOTYPE? Germany: Sachsen, Brambach im Vogtland Sohle in
altem Steinbruch, Granit. A. Spindler, 1908 (CANL 74962).
Stereocaulon tomentosum var. capitatum Lamb, J. Hattori Bot.
Lab. 43:3) '239) (1977 )%
HOLOTYPE. Argentina: Prov. Tucuman, Valle de Tafi, W. slope
of Cumbre Potrerillo. I.M. Lamb 5309, 1947 (CANL 63386).
Stereocaulon verruculigerum var. formosanum Asah., J. Jap.
Bot.23557).295> (1950)*
ISOLECTOTYPE. Taiwan: Rengechi. Y. Asahina 222, 1925 (CANL
63413).
= §. verruculigerum Hue var. verruculigerum
Stereocaulon vesuvianum f. verrucosum Lamb, J. Hattori Bot.
Labs! 43.59247°.(1577) &
ISOTYPE. United States: Alaska, Aleutian Islands, Attu. G. A.
Llano 1463 p.p., 1949 (CANL 77625). Poelt: Pl. Graecenses No.
246.
Stereocaulon vimineum Th. Fr., De Stereoc. & Pilophor.
Comment p. 13 (1857).
ISOTYPE? Mexico: Tiuzutlan. Liebmann 73, 1841 (CANL 75624).
= S. ramulosum (Sw.) Rausch.
Stereocaulon weberi Lamb, J. Hattori Bot. Lab. 43: 258
C1977 3%
ISOTYPES. Ecuador: Galapagos Islands, Isla Santa Cruz, summit
of Mount Crocker. W.A. Weber & J. Lanier, 15 April 1976 (CANL
58209, 77626). Weber: Lich. Exs. Colo. No. 494; Poelt: Pl.
Graecenses No. 247.
Sticta oregana Tuck., Bull. Torrey Bot. Club 5: 20 (1874).
ISOTYPE. United States: Oregon. E. Hall, 1871 (CANL 68763).
Reliq. Tuck. No. 90.
= Lobaria oregana (Tuck.) MUll. Arg.
Strangospora senecionis Lambinon & Vézda, Folia Geobot.
Phytotax., Praha 14: ' 206 (1979).
ISOTYPE. Zaire: Prov. Kivu, Birunga montes, in planitie
Rukumi montis ignivomi Karisimbi. Ad corticem et lignum
truncorum vetustorum. J. Lambinon 72/230, 22 January 1972
(CANL 73820). Vézda: Lich. Sel. Exs. No. 1657.
Sulcaria badia Brodo & D. Hawksw., Opera Bot. 42: 146 (1977).
ISOTYPE. United States: Oregon, Philomath. On old apple
trees. F.P. Sipe 669 (CANL 29671).
Note: This specimen is a portion of the same number
designated by Gyelnik as the type of Alectoria
pseudofuscescens Gyelnik. The holotype in BP, however, is
quite different. (fide Brodo & Hawksworth 1977. Opera Bot.
Aas 47) s
Sulcaria isidiifera Brodo, Mycotaxon 27: 115 (1986).
HOLOTYPE. United States: California, San Luis Obispo Co., Los
351
Osos Oaks State Reserve, Los Osos Valley Road. C. Bratt & J.
Larson 3871, 8 January 1984 (CANL 88639).
PARATYPE. United States: California. M.E. Hale 57816 (CANL
80150).
Tephromela elixii Kalb, Kalb: Lich. Neotr. Fasc. 12: 15
(1991).
ISOTYPE. Venezuela: Merida, Distr. Rangel, Paramo zwischen
der Laguna Mucubaji und dem Pico Mucufiuque, etwa 15 km SE von
Apertaderos. K. & A. Kalb, 16 August 1989 (CANL 103793).
Kal bsvLicn. aNeOtr..iNO. 522).
Tephromela muscicola Kalb, Kalb: Lich. Neotr. Fasc. 10: 15
(1991).
ISOTYPE. Ecuador: Tungurahua, Siidhang des Mt. Tungurahua,
nérdlich von Bafios. Uber Moosen in liickiger Vegetation. K. &
A. Kalb, 16 August 1987 (CANL 102736). Kalb: Lich. Neotr. No.
447.
Tephromela nashii Kalb, Kalb: Lich. Neotr. Fasc. 12: 16
(1991).
ISOTYPE. Mexico: Baja California, 10 km N El Rosario in Cafion
del Rosario. On small pebbles. K. & A. Kalb & T. Nash, 28
December 1990 (CANL 103794). Kalb: Lich. Neotr. No. 523.
Tephromela septentrionalis Hertel & Rambold, Bot. Jahrb.
Syst. 107: 495 (1985).
ISOTYPE. Greenland: W-Grénland, Gemeinde Umanak. Uber
eisenreichem Silikatgestein. J. Poelt & H. Ullrich, August
1983 (CANL 95897). Hertel: Lecideaceae Exs. No. 178.
Tephromela tropica Kalb, Kalb: Lich. Neotr. Fasc. 10: 15
(1988).
ISOTYPE. Kenya: Central Province, Nanyuki District, zwischen
Naco Moru und Nanyuki. K. Kalb & A. Schrégl, 18 August 1985
(CANL 102737). Kalb: Lich. Neotr. No. 448.
Tephromela velloziae Kalb, Kalb: Lich. Neotr. Fasc. 8: 16
(1984).
ISOTYPE. Brazil: Bahiz, Chapada Diamantina, Serra do
Tombador, etwa 1 km vor der Stadt Morro do ChapeG. K. Kalb,
18-20 July 1980 (CANL 89464). Kalb: Lich. Neotr. No. 349.
Thelotrema brasiliana Hale in Kalb, Kalb: Lich. Neotr. Fasc.
Py Sat 983 \,.
ISOPARATYPE. Brazil: Sao Paulo, Ilha de Sao Sebastido, etwa
130 km S6stlich von Sado Paulo. Westhang des Morro das Tacas.
In einem sehr feuchten und dunklen Regenwald an einem Fluss.
K. Kalb, 6 July 1979 (CANL 85936). Kalb: Lich. Neotr. No.
298.
Toninia arctica Timdal, Opera Bot. 110: 37 (1991).
HOLOTYPE. Canada: Northwest Territories, Franklin District,
Banks Island. G.W. Scotter 30146, 21-24 July 1979 (CANL
78540).
ISOTYPE. (CANL 78541).
352
PARATYPE. Canada: Yukon. G.W. Scotter 19501 (CANL 54403).
Toninia ruginosa var. andicola Lamb, Rhodora 56: 147 (1954).
HOLOTYPE. Argentina: Prov. Catamarca, Nevados de Anconquija,
Quebrada de los Cazadores. On soil. I.M. Lamb 5599, 24
November 1948 (CANL 6086).
= T. bullata (Meyen & Flotow) Zahlbr.
Toninia ruginosa subsp. pacifica Timdal, Opera Bot. 110: 90
(1991).
ISOTYPE. United States: California, Santa Barbara Co.,
Channel Islands, Santa Cruz Island, West end of the island.
On the rim of the "Trailer Barranca", in rock crevices. W.A.
Weber & C. Bratt, 8 January 1986. (CANL 97962). Weber: Lich.
Exs. Colo. No. 663.
Toninia tristis subsp. canadensis Timdal, Opera Bot. 110: 113
(1991).
HOLOTYPE. Canada: Northwest Territories, District of
Mackenzie, Dolomite Lake. G.W. Scotter 6097, 6 August 1965
(CANL 41669).
PARATYPES. Canada: British Columbia. I.M. Brodo 21333 (CANL
86793); Northwest Territories. G.W. Scotter 8283 & 8807 (CANL
43173 & 41670).
Toninia tristis subsp. pseudotabacina Timdal, Opera Bot. 110:
1a (1991)
PARATYPES. Cyprus: Larnaca. A. Vézda (CANL 99203). Vézda:
Lich. Sel. Exs. No. 2262; Spain: Prov. Larida. J. Poelt & A.
Vézda (CANL 89203).
Trapeliopsis hainanensis Hertel, Herzogia 5 (3-4): 460
(1981).
ISOTYPE. China: Prov. Guangdong, Insel Hainan. Bergwald-
Schutzgebiet im Gebirge Jian Fung Ling im Kreise Ledong. H.
Hertel, 23 May 1980 (CANL 77212). Hertel: Lecideaceae Exs.
Now -59:.
Trapeliopsis pseudogranulosa Coppins & P. James,
Lichenologist 16: 259 (1984).
ISOTYPE. Canada: British Columbia, Vancouver’ Island,
MacMillan Provincial Park on Rt. No. 4, 8 miles E of Alberni.
On Pseudotsuga. I.M. Brodo 7947, 5 June 1966 (CANL 53995).
Brodo: Lich. Canad. Exs. No 129.
Umbilicaria lambii Imsh., Bryologist 60: 232 (1957).
HOLOTYPE. Canada: British Columbia, Mt. Assiniboine
Provincial Park, Sunburst Lake. On a large rock. I.M. Lamb
6584, 3 August 1951 (CANL 66543).
Umbilicaria phaea Tuck., Lich. Calif. p. 15 (1866).
ISOTYPE. United States: California. On rocks of Pacific
Coast. H.N. Bolander 11, 1866 (CANL 68767). Reliq. Tuck. No.
94.
Usnea acromelana Stirton, Trans. & Proc. New Zealand Inst.
353
3035388 (1897)*
ISOTYPE. New Zealand: Christchurch, Canterbury, Selwyn Gorge.
"On trees" (saxicolous!). T.W.N. Beckett L.11, May 1894 (CANL
16943).
Usnea durietzii Mot., Lich. Gen. Usnea Stud. Monogr., Pars
Systems: 11s) 503 1(193 7)
ISOTYPE. Chile: Regio Magellanica, Insula Elisabetha. Ad saxa
erratica granitica. Lechler (CANL 17098).
Usnea eulychniae Follm., Nova Hedwigia 14: 261 (1967)
ISOTYPE. Chile: Prov. Antofagasta, zerstreut im Dornenwerk
von Sdaulenkakteen im Tornabenietum intricatae. I.A. Follmann-
Schrag & G. Follmann, September 1965 (CANL 64294). Follmann:
Lich. Exs7ysel. = No.=60:
Usnea kuehnemanrii Mot. ex Lamb, Farlowia 4: 467 (1955).
ISOTYPE. Argentina: Patagonia, Prov. Neuquén, Parque Nacional
Lanin, Lago Quillen. O. Kuhnemann 796, 1943 (CANL 104552).
Usnea neoguineensis Asah., J. Jap. Bot. 43: 496 (1968).
ISOTYPE. Papua New Guinea: Morobe District, Middle Creek
logging area, Bullolo. S. Kurokawa 5787, 11-12 November 1965
(CANL 34214). Kurokawa: Lich. Rar. Crit. Exs. No. 149.
Usnea neuropogonoides Mot., Lich. Gen. Usnea Stud. Monogr.,
pars System. I: 73° (1936).
ISOTYPE. Argentina: Rio Fosiles, in rupibus. P. Dusén, April
1905 (CANL 17188).
Usnea nidularis Asah., J. Jap. Bot. 44: 3 (1969).
ISOTYPE. Japan: Honshu, Prov. Iwaki, Matsukawa-ura, Sohma-
gun. On Pinus thunbergii. S. Kurokawa 58098-b, 18 July 1958
(CANL 65983). Kurokawa: Lich. Rar. Crit. Exs. No. 346.
Usnea pseudoceratina Mot., Folia Geobot. Phytotax. 10: 326
(1975).
ISOTYPE. Tanzania: Southern Highlands, Poroto Mountains, NW
end of Kitulo/Elton/Plateau. T. Pécs 6750/G, 10 August 1972
(CANL 57344).
Usnea tanzanica Mot., Folia Geobot. Phytotax., Praha 10: 327
(1975).
ISOTYPE. Tanzania: Prov. Mbeya, in monte ignivomo Rungwe. Ad
ramulos arborum in regione subalpina. T. Pécs 6507, 21
January 1972 (CANL 57292). Vézda: Lich. Sel. Exs. No. 1322.
Verrucaria bagliettoi Servit, Ann. Mus. Civico Storia Nat.
Giacomo Doria 64: 53 (1950).
ISOTYPE. Italy: Genova, Val Bisagno, Rupicola loco Fullo.
Sbarbaro, November 1950 (CANL 692).
Verrucaria bisagnoensis Servit, Sborn. Nar. Mus. v Praze,
Rada B, Pfrir. Védy. 9: 16 (1949).
ISOTYPE. Italy: Genova, Val Bisagno, Prato. Sbarbaro, April
1949 (CANL 693).
354
Verrucaria boccana Servit, Sborn. Nar. Mus. v Praze, Rada B,
Prir. Védy. 9: 17 (1949).
ISOTYPE. Italy: Dalmatia meridionalis, Hercegnovi 40 nm,
Silic. M. Servit, 1929 (CANL 694).
Verrucaria contardinis Servit, Ann. Mus. Civico Storia Nat.
Giacomo Doria 66: 236 (1953).
ISOTYPE. Italy: Friuli, Zomeais, prope Tarcento. A. Contardo,
25 December 1951 (CANL 711).
Verrucaria diplotommoides Servit, Preslia 24: 355 (1952).
ISOTYPE. Italy: Genova, Staglieno. Sbarbaro, 30 March 1951
(CANL 718).
Verrucaria durietzii Lamb, Lilloa 14: 205 (1948).
HOLOTYPE. New Zealand: Auckland Islands, Port Ross, innermost
small peninsula in Laurie Harbour. G.E. Du RietzZ 2225 b:1, 27
March 1927 (CANL 721).
PARATYPE. New Zealand. G.E. & G. Du Rietz 2070:2 (CANL 724).
Verrucaria durietzii f. rhabdota Lamb, Lilloa 14: 206 (1948).
HOLOTYPE. New Zealand: E. Falkland Islands, Berkeley Sound,
Port Louis. On seashore rocks. I.M. Lamb 2939, 9 February
1946 (CANL 720).
PARATYPE. New Zealand. I.M. Lamb 2877, 29 January 1946 (CANL
723
Verrucaria fuscella f. omblensis Servit, Stud. Bot. Cech. 7:
59 (1946).
ISOTYPE. Italy: Dalmatia, Ragusa, Ombla, Gionchello. Latzel
104, 1908 (CANL 802).
= V. omblensis (Servit) Servit
Verrucaria gorzegnoensis Servit, Webbia 8: 413 (1952).
ISOTYPE. Italy: Cuneo, Gorzehno, le Langhe. Sbarbaro,
September 1951 (CANL 744).
Verrucaria haeyrenii ("hayrénii") Erichsen, Memoranda Soc.
Faunavhl. Fenn .l2ice Se (1937 ) i.
ISOTYPE. Finland: Nyland, Ekends, Tvdrminne, Langskar. E.
Hayrén & C.F.E. Erichsen, 29 June 1933 (CANL 747).
Verrucaria imperfecta Servit, Preslia 24: 359 (1952).
ISOTYPE. Italy: Genova, Val Risagno, Ligorna. Sbarbaro, 21
September 1950 (CANL 749).
Verrucaria incompta Sersit, Ann. Mus. Civico Storia Nat.
Giacomo Doria 66: 237 (1953).
ISOTYPE. Italy: Savona, Spotorno, Rupicola loco Lajolo.
Sbarbarc, January 1952 (CANL 752).
Verrucaria lacustris Lamb, Farlowia 4: 424 (1955).
HOLOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Verde near
Futalaufquen. On stone. I.M. Lamb 5884, 1 February 1950 (CANL
760).
355
Verrucaria langhensis Servit, Ann. Mus. Civico Storia Nat.
Giacomo Doria 64: 48 (1950).
ISOTYPE. Italy: Gorzegno, Le Langhe, Piemonte. Sbarbaro,
September 1947 (CANL 764).
Verrucaria latebrosoides Servit, Sborn. Nar. Mus. v Praze,
Rada B, Piir. Védy. 9: 31 (1949).
SYNTYPE. Italy: Genova, Val Bisagno, Ligorna. Sbarbaro, 23
March 1950 (CANL 766).
Verrucaria ornata Servit, Stud. Bot. Cech. 11: 119 (1950).
ISOTYPE. Italy: Genova, Mt. Fasce. Sbarbaro, 1947 (CANL 803).
Verrucaria pseudomacrostoma Servit, Webbia 8: 415 (1952).
ISOTYPE. Italy: Cuneo, Langhe, Gorregno, 400 mt. Cortemilia.
Sbarbaro, September 1951 (CANL 809).
Verrucaria putnae Servit, Preslia 24: 374 (1952).
ISOTYPE. Romania: Putna, Sovejanka. Cretzoiu, 1935 (CANL
613).
Verrucaria rapallensis Servit, Stud. Bot. Cech. 11: 121
(1950).
ISOTYPE. Italy: Genova, Rapalleo, Montallegro 500 mt.
rupicola. Sbarbaro, October 1949 (CANL 814).
Verrucaria santensis Nyl., Acta Soc. Sci. Fenn. 7: 489
(1863).
ISOLECTOTYPE. United States: South Carolina, Santee Canal.
H.W. Ravenel, 1850 (CANL 23286).
= Pyrenula santensis (Nyl.) Miill. Arg.
Verrucaria savonensis Servit, Webbia 8: 416 (1952).
ISOTYPE. Italy: Savona, Albisola marina, loco Seitun.
Sbarbaro, August 1951 (CANL 822).
vézdaea rheocarpa Poelt & Débbeler, Bot. Jahrb. Syst. 96: 347
(1975).
ISOTYPE. Austria: sitidliches Burgenland. Uber verschiedenen
Moosen. J. Poelt & P. Ddébbeler, 17 March 1974 (CANL 63545).
Poelt: Lich. Alpium No. 293.
Wawea fruticulosa Henssen & Kantvilas, Lichenologist 17: 86
(1985).
ISOTYPE. Australia: Tasmania, Cradle Mountain National Park.
On burned trunk of Nothofagus cunninghamii. W.A. Weber & D.
McVean, 22 February 1968 (CANL 52202). Weber: Lich. Exs.
COLO. INO. 452).
Xanthoparmelia cylindriloba Knox, J. South African Bot. 49:
147 (1983).
ISOTYPE. Kenya: Regio montis Kenya, in parte superiore vallis
Teleki. Ad terram glacie destructam. O. Hedberg, 27 July 1948
(CANL 93769). Vézda: Lich. Sel. Exs. No. 2063.
Xanthoparmelia lipochlorochroa Hale & Elix, Mycotaxon 34: 551
356
(1989).
ISOTYPE. United States: Wyoming, Lincoln Co., 15 mi. S of
Kemmerer, Red Desert area. On sandy soil. B.J. Buckingham S
22155, 26 August 1959 (CANL 20819). Weber: Lich. Exs. Colo.
NO. ‘29.
Xanthoparmelia mollis Hale, Mycotaxon 27: 585 (1986).
ISOTYPE. South Africa: Natal. Port Shepstone District, Oribi
Gorge Nature Reserve. On exposed sandstone outcrops. M.E.
Hale 74043, 3 February 1986 (CANL 99574). Almborn: Lich.
African, iNos 1237
Xanthopeltis rupicola R. Sant., Svensk Bot. Tidskr. 43: 560
(1949).
ISOTYPE. Chile: Prov. Santiago, San José. On rocks. R.
Santesson 2420, 12 June 1940 (CANL 18087).
Xanthoria filsonii Elix, Elix: Lich. Austral. Exs. Fasc. 7
Nove 17491988):
ISOTYPE. Australia: Victoria, Hopetoun, Patchewollock Road,
5 km NE of Hopetoun. On Rhagodia. J.A. Curnow & H. Lepp 1422,
14:-March 1987. * (GANL (98903),. Elix: Lich: Austral. Exs. No.
aes
Xanthoria polycarpa var. maritima Lamb, Rep. (Annual) Natl.
Musi; Canada,, 1992-54. Bull 213238 306 mo 54 Fr
HOLOTYPE. Canada: Nova Scotia, Victoria Co., Cape Breton
Island, Ingonish, Middle Head. On granite rocks. I.M. Lamb
6952, 29 July 1952 (CANL 18084).
Zahlbrucknerella patagonica Henssen, Lichenologist 9: 41
(1977).
ISOTYPE. Argentina: Prov. Santa Cruz, Lago Argentino,
Calafate, Cuevas de Hualichu. Sickerwasserflachen auf
Schieferfels in Uferndhe. A. Henssen & G. Vobis 24525a, 18
December 1973 (CANL 102535). Henssen: Lich. Cyanoph. Fungi
Sax see NOs) 6 Us
ISOPARATYPE. Argentina. I.M. Lamb 5858 p.p. (CANL 27339).
ACKNOWLEDGEMENTS
To thank (Drow. Mee Brodo and Drow rk. Kya relandm.or
valuable suggestions and comments and for corrections to the
manuscript, Dr. T. Ahti for critical review of this paper. I
also thank Miss L. Ley for checking the abbreviations of
periodicals.
357
REFERENCES
Bridson, ei) ey 399019, Botanico-Periodicum-
Huntianum/Supplementum. Carnegie-Mellon University,
Pittsburgh.
Egan, R.S., 1987. A fifth checklist of the lichen-forming,
lichenicolous and allied fungi of the Continental United
Stated and Canada. Bryologist 90: 77-173.
Egan, R.S., 1989. Changes to the "Fifth Checklist of the
Lichen-Forming, Lichenicolous and Allied Fungi of the
Continental United States and Canada." Edition I.
Bryologist 92: 68-72.
Egan, R.S., 1990. Changes to the "Fifth Checklist of the
Lichen-Forming, Lichenicolous and Allied Fungi of the
Continental United Stated and Canada." Edition II.
Bryologist 93: 211-219.
Egan, R.S., 1991. Changes to the "Fifth Checklist of the
Lichen-Forming, Lichenicolous and Allied Fungi of the
Continental United States and Canada." Edition III.
Bryologist 94: 396-400.
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MY COTAXON
Volume XLVI, pp. 359-366 April-June 1993
NEOSARTORYA PRIMULINA, A NEW SPECIES OF FOOD-
BORNE ASCOMYCETES
Shun-ichi UDAGAWA!, Noritsuna TOYAZAKI¢
and
Haruo TSUBOUCHI>
1 Nodai Research Institute, Tokyo University
of Agriculture, 1-1-1, Sakuragaoka, Setagaya-
ku, Tokyo 156, Japan
2 Public Health Research Institute of Kobe
City, 4-6, Minatojimanaka-machi, Chuo-ku,
Kobe 650, Japan |
3 Nagoya City Health Research Institute, 1-11,
Hagiyama-cho, Mizuho-ku, Nagoya 467, Japan
ABSTRACT
A new species of Neosartorya, N. primulina
Udagawa, Toyazaki et Tsubouchi, isolated from a
canned oolong tea beverage in Japan, is de-
scribed and illustrated. This fungus is char-
acterized by its restricted growth on Czapek
agar, chalky-buff ascomata, and lenticular
ascospores with a very irregular ornamentation
composed of several narrow crests and verrucose
hemispheres.
Recently some widely-distributed species of Neosarto-
rya have been incriminated as spoilage agents in commer-
cially processed fruit products (Kavanagh et al., 1963;
Beuchat, 1986; Conner and Beuchat, 1987: Scott and
Bernard, 1987; Gomez et al., 1989; Nielsen et al., 1989;
Samson, 1989; Splittstoesser and Churey, 1989; Samson et
al., 1992). In an earlier paper (Udagawa et al., 1991),
Neosartorya hiratsukae Udagawa, ITsubouchi et Horie was
reported as the new causal agent of spoilage of pasteur-
ized aloe beverage in Japan. A variety of commercial pas-—
teurized beverages: which were undergoing spoilage were
360
selectively surveyed for further isolation of heat resist-
ants. Pasteurized beverages (packaged in cans) by com-
mercial processors for fungal isolation. Fungal mycelia
were removed from the beverage by straining through a
sterile membrane filter by aseptically transferred to
chloramphenicol (100 ug/1)-potato dextrose agar.
During the course of this survey, several heat re-
sistant fungi were encountered. Among them one isolate
representing a hitherto undescribed species of Neosartorya
(Malloch and Cain, 1972; Kozakiewicz, 1989; Peterson, 1992)
was found which is herein described and illustrated,
Neosartorya primulina Udagawa, Toyazaki et Tsubouchi,
Sp. nov. GiGicge hee
St. Anam. Aspergillus primulinus Udagawa, Toyazaki et
Tsubouchi, anam. sp. nov.
Coloniae in agaro Czapekii paulo restrictae, aliquan-
tum floccosae, plus minusve radiatim sulcatae, laxae,
flavo-albae; conidiogenesis sparsa; reversum incoloratum
vel pallide flavum. Coloniae in "Czapek yeast extract
agar (CYA)" celeriter crescentes, velutinae, conspicue
sulcatae, saepe zonatae, ex coacto mycelio basali tenuiter
constantes, numerosis ascomatibus formantes, granulares,
flavo-albae vel pallide flavae vel primulinae; conidio-
genesis varians, viridi-grisea vel avellanea; reversum
pallide flavum vel rubro-brunneum vel brunneo-aurantiacum.
Coloniae in agaro maltoso effusae, floccosae, parum zon-
atae, tenues, laxae, abundantibus ascomatibus formantes,
pallide flavae; conidiogenesis inconspicua; reversum
incoloratum vel pallide flavum.
Ascomata non-ostiolata, superficialia, dispersa,
cretacea vel bubalina, globosa vel subglobosa, 100-250 um
diam, hyphis brunneis laxe intricatis circumdata, lente
maturescentia; peridium membranaceum, ‘textura epidermoi-
dea’ et "textura angularis'; stratum externum ex cellulis
brunneis, irregularibus, incrassatis, 5-/.5 um latis com-
positum. Asci octospori, brevi-catenati, subglobosi vel
plus minusve pyriformes, 12-15(-17.5) x 11-13(-15) um,
evanescentes. Ascosporae hyalinae, lenticulares, sine
cristis 4.5-5(-5.5) x 3.8-5 um, cristis numerosis angusta-
tis irregulatim ornatae.
Capitula conidica parva, brunneo-grisea, laxe radian-
tia; conidiophora ex mycelio basali vel interdum hyphis
aeriis ascendentes; stipites brunneoli, 200-600 x 2.5-5
(-6) um, leves et incrassati, saepe septati; vesiculae
brunneae, ampulliformes, 7.5-16 um diam. Aspergilla uni-
serialia; phialides cylindricae, 5-7.5 x 2-2.5 um, in sum-
361
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362
ma 1/2-2/3 vesicula insidentes. Conidia hyalina, globosa
vel subglobosa vel parum ovoidea, 2-3(-4.5) x 2-2.5 um,
levia.
Holotypus SUM 3014, colonia exsiccata in cultura ex
aqua decocta thea, Okinawa in Japonia, 22.vii.1991, iso-
lata et ea in collectione fungorum Musei et Instituti
Historiae Naturalis Chiba (CBM) conservata.
Etymology: from Latin, primulinus=primrose, referring
to thevcorvonveco tan:
Colonies on Czapek agar growing rather restrictedly,
attaining a diameter of 27 mm within 7 days at 25°C,
somewhat floccose, more or less furrowed in a radial pat-
tern, loose-textured, yellowish white (M. 4A2 after
Kornerup and Wanscher, 1978); conidiogenesis sparse, in—
conspicuous; reverse uncolored to pale yellow (M. 4A3) or
Straw (Rayner, 1970). Colonies on CYA growing rapidly,
attaining a diameter of 46 mm within 7 days at 25°C, or
85 mm within 7 days at 37°C, velvety, deeply furrowed,
often zonate, consisting of a thin basal felt intermixed
with numerous ascomata in a granular appearance at 25°C,
yellowish white (M. 4A2) to pale yellow (M. 3A3-4A3) or
Primrose (Rayner); conidiogenesis limited or profuse at
37°C, in the latter case greenish gray to brownish orange
(M. 29D2-6C4) or Hazel (Rayner); reverse light yellow to
reddish brown (M. 4A4-8E5) or Dark Brick (Rayner). Colo-
nies on malt extract agar (MEA) spreading broadly, attain-
ing a diameter of 54-56 mm within 7 days at 25°C, or 85
mm within 7 days at 37°C, floccose, plane or slightly zon-
ate, thin, loose-textured, characterized by abundant asco-
mata in granular appearance, pale yellow (M. 4A3) or Buff
(Rayner); conidiogenesis moderate at 37°C but not suffi-
ciently produced to influence the colony appearance; re-
verse uncolored to pale yellow (M. 4A2-3) or Pale Luteous
(Rayner).
Ascomata non-ostiolate, superficial, scattered,
chalky to buff, globose to subglobose, 100-250 um in diam,
covered loosely with brownish encrusted hyphae, maturing
within 21 days at 25°C; peridium 10-12.5 um thick, membra-
naceous, textura epidermoidea, two-layered; outer layer
composed of brownish, irregular-shaped, thick-walled cells
measuring 5-/.5 wm wide; inner layer of hyaline, thin,
Figure 2. Neosartorya primulina, SUM 3014.
A. Ascosporesu( SEM) ox co00eambe Aseria oo:
C. Conidia, x 1300. D, E. Aspergilla, x 1000 and
x 1300, respectively.
363
364
Ascomatal initials developing as short branches of hyphae
with large curled tips measuring ca. 12.5 um in diam. Asci
8-spored, produced in short chains, subglobose to more or
less pyriform, 12-15(-17.5) x 11-13(-15) um, evanescent at
maturity. Ascospores hyaline, lenticular, spore body 4, 5-
5(-5.5) x 3.8-5 um, provided with several narrow crests
measuring ca. 0.5 um wide and with irregularly verrucose
hemispheres. Conidial heads intermixed with ascomata,
small, brownish gray, loosely radiate; conidiophores aris-—
ing from the basal mycelium or sometimes from the aerial
hyphae, stipes brownish, 200-600 x 2.5-5(-6) um, smooth
and thick-walled, often several times septate; vesicles
brownish, flask-shaped, 7.5-16 um in diam. Aspergilla uni-
seriate, phialides cylindric, 5-/.5 x 2-2.5 um, covering:
the upper half to two-thirds of the vesicle. Conidia hya-
line, globose to subglobose or slightly ovoidal, 2-3(-4.5)
x 2-2.5 um, smooth-walled. Maximum growth temperature:
47°C. Pasteurization» atyee.C for 15 min. 1s effective.
Specimen examined: SUM 3014 (holotype), a dried cul-
ture of an isolate from a canned oolong tea beverage, Oki-
nawa-Pref., Japan, 22.vii.1991. The specimen studied and
living culture derived from the type are preserved in the
Natural History Museum and Institute, Chiba (CBM), Japan.
Neosartorya primulina can be readily recognized by its
chalky-buff colored ascomata, short-stipitate initials of
the ascomata, and morphology of the ascospores. The scann-
ing electron micrograph (SEM) of the ascospores provided a
much clearer differentiation between N. primulina and all
other members of Neosartorya (Samson et al., 1990, 1992);
the ascospore ornamentation in this species is character-
ized by several narrow crests in the equatorial area and
irregularly verrucose hemispheres (Fig. 2). Neosartorya
hiratsukae somewhat resembles this species in the restrict-
ed growth on Czapek agar and in the formation of pale yel-
low ascomata, but differs in having ascospores with convex
walls bearing numerous closely anastomosing ridges arrang-
ed in a fine reticulate ornamentation(Udagawa et al.,1991).
Acknowledgement: We thank Prof. D. Malloch of the Univer-
sity of Toronto for reviewing this paper.
Literature cited
Beuchat, L. R. 1986. Extraordinary heat resistance of
Talaromyces flavus and Neosartorya fischeri asco-
SHOP esmenicru le prROGUCUSiwm Us Food! SGT, canine LoUG-
15708
365
Conner, D. E. and Beuchat, L. R. 1987. Heat resistance of
ascospores of Neosartorya fischeri as affected by
sporulation and heating medium. Intl. J. Food
Microbiol., 4: 303-312.
Gonezeaen Mae DUStagereure sander lug, imu.) 1989s Sef fect
of the post-dry heat treatment temperature on the
recovery of ascospores of Neosartorya fischeri.
Lett. Appl. Microbiol.,.8: 59-62.
Kavanagh, J., Larchet, N. and Stuart, M. 1963. Occurrence
of a heat resistant species of Aspergillus in canned
strawberries. Nature (Lond. ), 198: 1322.
Kornerup, A. and Wanscher, J. H. 1978. Methuen Handbook
of Colour. 3rd ed. Eyre Methuen, London. 252p.
Kozakiewicz, Z. 1989. Aspergillus species on stored pro-
ducts. CMI Mycological Papers No. 161: 1-188.
Malloch, D. and Cain, R. F. 1972. The Trichocomataceae:
Ascomycetes with Aspergillus, Paecilomyces, and Peni-
cillium imperfect states. Can. J. Bot., 50: 2613-
cocae
i ciliscntmeny Seubeuchaues ark mranGeaprisvVadscds"U, 919097
Growth and fumitremorgin production by Neosartorya
fischeri as affected by food preservatives and organic
acids. J. Appl. Bacteriol., 66: 197-207.
Peterson, S. W. 1992. Neosartorya pseudofischeri sp. nov.
and its relationship to other species in Aspergiilus
section Fumigati. Mycol. Res., 96: 547-554,
Rayner, R. W. 1970. A Mycological Colour Chart. Common-
wealth Mycological Institute, Kew and the British
Mycological Society.
Samson, R. A. 1989. Filamentous fungi in food and feed.
Je App ls Bacteriol). Symp. supp ls .t/07s 275-355.
Samson, R. A., Nielsen, P. V. and Frisvad, J. C. 1990. The
genus Neosartorya: Differentiation by scanning elect-
ron microscopy and mycotoxin profiles. In: Modern
Concepts in Penicillium and Aspergillus Classifica-
Ci OncO mrSamscOnmenk een emander tbCend sete ee 405-401,
Plenum Press, New York.
SANSOM mane MA em HOCKING ee AU sence Camu eau, MONGEN UNC aiee ns
(ed. ) 1992. Modern Methods in Food Mycology.
Elsevier, Amsterdam. 388p.
Scott, V. N. and Bernard, D. T. 1987. Heat resistance of
Talaromyces flavus and Neosartorya fischeri isolated
from commercial fruit juices. J. Food Prot., 50: 18-
(A>.
Solittstoesser, 0: Fovand Churey,- J. JU, .1989." Effect of
low concentration of sorbic acid on the heat resist-
ance and viable recovery of Neosartorya fischeri asco-
366
spores. J. Food Prot., 52: 821-822.
Udagawa, S., Tsubouchi, H. and Horie, Y. 1991. Neosarto-
rya hiratsukae, a new species of food-borne Asco-
mycetes. Trans. Mycol. Soce-Wapany. ocimes-49.
MY COTAXON
Volume XLVI, pp. 367-377 April-June 1993
STUDIES ON BOLETUS SECTION LURIDI
Re. TREU
Virginia Polytechnic Institute and State University,
Department of Biology, Blacksburg, Virginia, 24061
SUMMARY
Four species of Boletus section Luridi described by Smith
and Thiers have been studied: Boletus pseudo-olivaceus, B.
roseobadius, B. rufocinnamomeus, and B. subluridellus.
B. pseudo-olivaceus can be easily differentiated from the
other species by its weakly but distinctly amyloid trama in
stipe and pileus. There are no major microscopical differ-
ences between B. subluridellus, B. rufocinnamomeus, and B.
roseobadius.
INTRODUCTION
The section Luridi Fr. is a relatively well defined
group in the genus Boletus Fr. Singer (1986) characterized
the section by its Boletus habit, the narrow pores, which
are often discolorous and red, and by the occasional
presence of toxins in the fruitbodies. The sporocarps turn
immediately blue on bruising or cutting. Smith and Thiers
(1971), who interpreted the group as a subsection within
the section Boletus, have increased our knowledge of the
group considerably; they also described several new species
from Michigan.
A recently found collection from Virginia revealed
however, that species delimitation within this group is
still problematic; a number of characters considered to be
important for species delimitation showed variation within
the collection. Smith and Thiers (1971) described a
"Spatter pattern" for the distribution of different
characters within this group and suspected a gene exchange
between some taxa, such as B. subvelutipes Peck and B.
pseudo-olivaceus Smith and Thiers, two species growing in
the same habitat. Therefore, a reevaluation of characters
seems to be necessary. As a first step the collection from
Virginia and specimens of four different species which are
closely related have been studied here.
368
MATERIALS AND METHODS
Microscopical observations were made in KOH (10%)
unless stated otherwise, but the cuticle was observed in
Melzer's reagent. A drawing tube was used for the illustra-
tions. At least 15 spores per collection were measured and
the mean dimensions D" for the spores were calculated; the
length/width ratio (Q) was obtained for each of the spore
measurements and a mean value (Q,) was calculated for each
of the species.
Detailed descriptions of the macroscopic characters of
each taxon were presented by Smith and Thiers (1971).
Color descriptions follow Kornerup & Wanscher (1967).
Official colors are in quotes.
The terminology of cystidia has been adopted from
Singer (1986) and Wolfe (1991).
RESULTS
BoLeeus wR 316
Figs 1521-25
Pileus 4-14 cm, convex, surface tomentose to almost
velvety, »color.darks"olive’ brown" (tou"raw umber"). "soot
brown", or "coffee", (4 F4-8; 5 F5-8), a few fruitbodies
with rose-red areas. Tubes adnexed, up to 1 cm thick,
yellow, but rapidly staining blue when cut. Pores round,
small, “lake red", "Pompeian red", "dark red" until old age
(9 C7-8, 10 C7-8), staining greenish blue when injured.
Spore print "bronze", "mustard brown", "linoleum brown" (5
E5-7).
Stipe clavate or equal, solid, 2.5-9 cm long, 1.5-3.5
cm thick, orange-yellow, dark red toward the base, bruising
instantly blue. In some specimens a fine red reticulation
was observed on the very tip of the stipe.
Context of the whole fruitbody after cutting instantly
changing to dark blue.
Chemical reactions of (blued) pileus context: KOH
orange; NH,OH yellow, FeSO, yellow. Chemical reactions on
pileus supiace:s KOH (—)/7)NH,OH i>) FeSO .a(—)e
Spores 10:8-13.5) x 3%6-5/4 pm; D" = 12.1% 4.4. pm780
= 92.353 537 0 J= 2.8 ;etusoid,, yellow to, brownishsyellowiin
KOH; spores from deposit 11.7-14.4 x 3.6-5 um, pale yellow-
brown in KOH; spores in Melzer's reagent 10.4-12.6 x 3.6-
4.5 um, occasionally thick-walled, pale yellow-brown.
Basidia 26-41 x 8-10 um, 4-spored, occasionally 2-spored,
no basal clamps, hyaline in KOH and Melzer's reagent, a few
with a yellow, plasmatic pigment. Pleurocystidia 24-52 x
4.7-8 wm, fusiform. Pseudocystidia 19-31 x 3.7-5.8 wun,
fusiform. All septa without clamps.
Cuticle a trichoderm or occasionally a cutis, 180-190
um thick; trichodermial end cells connected at their tips
and forming fascicles. Trichodermial hyphae 3.5-4.1 pm
369
thick, with deep red walls and occasionally brown contents
in Melzer's reagent; yellowish, pigment plasmatic, some-
times with granular contents in KOH. Context below tricho-
derm a tangle of hyaline hyphae, 3.5-5.9 wm thick, with
granular, yellow-brown debris between hyphae; occasionally
some hyphae have peculiar wall thickenings up to 1.5 um.
Oleiferous hyphae can be found occasionally (diam. 5 pum).
Stipe dermatocystidia of two types: first type 21-26
x 4.2-7.4 um clavate, similar to probasidia. Second type
(less frequent) 19-110 x 4.1-8 wm, fusiform. Scattered
basidia can be found all along the stipe cortex, 23-35 x 8-
10 um, 2- or 4-spored, with yellow content in KOH, yellow-
brown to dark brown in Melzer's reagent. Tramal hyphae
inamyloid.
Comments. This collection seems to be close to B.
subluridellus although the Q value for the spores is some-
what higher for this collection (2.8 here versus 2.6 in the
holotype of B. subluridellus). A number of characters shows
high variation within the collection:
The long and setiform caulocystidia which were
regarded by Smith and Thiers (1971) as an important
character to delimit the group around B. subluridellus
could be found only sporadically in some but not all
fruitbodies of this collection. Therefore their presence
seems to vary within at least this species.
The structure of the cuticle is also variable: usually
it consists of conical fascicles of trichodermial hyphae
but in the same collection it can be a cutis.
Collection examined:
Virginia, Montgomery Co., near Pandapas Pond, under oak and
white pine, 9 Aug 1991, R. Treu (TR 1316), Massey Herbarium
(VPI).
Boletus pseudo-olivaceus Smith & Thiers. 1971. The boletes
OLPMa Chigan awe 5s:
Firgss [=5
Spores 12.2-15.3 x 5.4-6.3" pm; DY = '141.%'°5.9 pm?'OQ
=2 3726.07, Or = 2.45) fusoid, often thick-walled}(0.5=-1 um),
in KOH yellow; spores in Melzer's reagent 9.9-15.3 x 5-5.9
um, thick-walled (0.5-1 wm), pale brownish yellow. In
Melzer's reagent many spores show a minute, indistinct germ
pore. Basidia 31-39 x 9-13 wm, four-spored or two-spored,
hyaline or yellow. Pleurocystidia 37-41 x 6-7 um, fusiform.
Pseudocystidia 26-29 x 6-7 um, clavate, with golden yellow
pigmentation (plasmatic). All septa without clamps.
Cuticle a cutis, 350-450 um thick, with many oleifer-
ous hyphae; hyphae 2.3-6.3 um thick, hyphal walls not red
in Melzer's reagent; oleiferous hyphae with yellow, homoge-
neous content.
Stipe dermatocystidia 29-63 x 12-14 um, fusiform or
370
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‘eTptsed ~ ‘sezods T ‘*“SneoeATTO-opnes
“w7#O0T = saeq eTeos
-etptysAoopnesd €
SnjeTOd G-T °*shTa
371
clavate, yellow or brownish yellow. Occasionally yellow
caulobasidia can be found among the cystidia. Tramal hyphae
of stipe as well as tramal hyphae of pileus slightly
amyloid, purple to grayish in Melzer's reagent. Yellow-
orange tomentum at the base of the stipe consisting of
bundles of parallel hyphae, 1.8-3.6 wm thick, yellowish in
KOH, brownish and some hyphae with dark brown contents in
Melzer's reagent.
Comments. B. pseudo-olivaceus is easily distinguished
from the other species by a weak, but distinct amyloid
reaction of the hyphae in both the pileus and the stipe
trama. This reaction is persistent and should not be
confused with the "fleeting-amyloid" reaction described by
Smith and Thiers (1971) for some boletes. The spores of B.
pseudo-olivaceus also have a lower Q, value (2.4) than the
other species (2.6-2.7).
Collections examined:
Michigan, Emmet Co., Mackinaw City Hardwoods, scat-
tered at edge of hardwood forest, 12 Jul 1967, A.H.
Smith 74510*- holotype, MICH.
Michigan; Charlevoix) Co.) Eastidordan; ,12%Jul) 1967,
James Bennet (A.H. Smith 74511), MICH.
Boletus roseobadius Smith & Thiers. 1971. The boletes of
Michigan. Ps )354.
Figs. 6-10
Spores 13.1-16.2 x 4.5-6.3 pm; DY = 14.5 x 5.5 pm; Q
= 2.4-3; Q = 2.7; fusoid, sometimes thick-walled, yellow-
ish-brown to yellow in KOH; spores in Melzer's reagent
12.2-16.7 x 4.5-6.3 wm, yellowish-brown to brown. Spores on
the stipe cortex sometimes dextrinoid and broader than
hymenial spores. Basidia 25-34 x 8-12 wm, four-spored or
two-spored, hyaline or sometimes with yellow, homogenous
or granular, plasmatic pigment. Pleurocystidia 24-62 x 6-10
um, fusiform, mostly with apical neck. Pseudocystidia 21-34
x 5-6 um, fusiform or clavate, with golden yellow pigmenta-
tion (plasmatic), along the tube edges; occasionally a few
pseudocystidia with size and shape of the pleurocystidia,
but with yellow pigment, on the tube walls. All septa
without clamps.
Cuticle a cutis, 130-230 wm thick; hyphae 2.7-5.4 um
thick, hyphal walls red in Melzer's reagent; some hyphae
with yellow, granular content.
Stipe dermatocystidia 24-80 x 5-11 wm, fusiform or
cylindric sometimes with an apical neck, hyaline or brown.
Occasionally 3- or 4-spored brown basidia can be found
among the cystidia. Tramal hyphae inamyloid, in stipe base
sometimes with red walls.
Comments. It is almost impossible to distinguish this
372
10
i Pre
Figs. 6-10 Boletus roseobadius. 6 Spores. 7 Basidia
8 Pleurocystidia. 9 Pseudocystidia. 10 Stipe dermatocystidia.
Figs. 11-15 Boletus rufocinnamomeus. 11 Spores. 12 Basidia
13. Pleurocystidia. 14 Pseudocystidia. 15 Stipe dermato-
cystidia. Scale bars = 10ysm.
373
species from B. subluridellus on microscopic characters. A
very distinctive feature of these species is the deep red
color of the walls of many cuticular hyphae in Melzer's
reagent. Smith and Thiers (1971) characterized the species
by the size of the pleurocystidia, the red (in Melzer's
reagent) cuticular hyphae, and the brown cortex elements
(in KOH). However, there is no differences in the size of
the pleurocystidia (24-62 um in B. roseobadius, 34-50 um in
B. subluridellus). Smith and Thiers (1971) described B.
roseobadius as having a whitish or pallid flesh in compari-
son the bright yellow flesh in B. subluridellus. The
interpretation of this feature might be difficult because
of the instant color change of the flesh to deep blue.
Collections examined:
Michigan, Washtenaw Co., solitary under low hardwoods,
20 Sep 1965, F. V. Hoseney (A.H. Smith 72668 - holo-
type), MICH.
Michigan, Washtenaw Co., Pinckney State Recreation
Areas) 258 Auge 9 ish A. HeSmatn $8453:-7:, IMiCH:
Boletus rufocinnamomeus Smith & Thiers. 1971. The boletes
OTeMVchiganiiPses5 Gy
Figss214-15
Spores 13.2-16.2 x 5.1-6.3 pm; DY = 14.4 x 5.4 pm; Q
=e 4-319; 0.) = 92,75) Lusoid; amostily, thin-walled,, pale’to
golden yellow; spores in Melzer's reagent 12.6-17.1 x 4.5-
6.3 um, yellowish brown. Basidia 30-34 x 10-12 um, four-
spored, occasionally two-spored, hyaline. Pleurocystidia
24-40 x 5-9 um, fusiform with apical neck. Pseudocystidia
25-34 x 4-6 pum, fusiform, sometimes with a neck, with
golden yellow pigmentation (plasmatic), along the tube
edges, sometimes a few on the tube walls. All septa without
clamps.
Cuticle a trichodermium, 180-200 um thick, formed of
conical fascicles of trichodermial hyphae with brown, often
granular content in Melzer's reagent; occasionally some of
the hyphae below the fascicles with weakly red walls in
Melzer's reagent. Hyphae forming the fascicles 1.8-3.6 um
thick, hyphae below 2.7-6.3 um.
Stipe dermatocystidia 24-100 x 4-14 um, fusiform, with
or without a neck, occasionally with a secondary septum,
hyaline or golden yellow or brownish. Tramal hyphae
inamyloid.
Collections examined:
Michigan, Cheboygan Co., Burt Lake, hardwoods, 11 Aug
1969, A.H. Smith 77831 - holotype, MICH.
Michigan, Cheboygan Co., Burt Lake, Colonial Point
Hardwoods, 7 Aug 1969, A.H. Smith 77746, MICH.
374
Boletus subluridellus Smith & Thiers. 1971. The boletes of
Michigan. P. 349.
Figs. 16-20
Spores 10.6-15.3 x 4.5-5.4 um; DY = 12.8 x 4.9 um; Q
= 2.2-2.9; Q. = 2.6; fusoid, occasionally slightly thick-
walled, brownish yellow in KOH; spores in Melzer's reagent
11.7-14.4 x 4.5-5.9 um, yellow-brown. Spores on stipe
cortex frequently with golden yellow pigmentation. Basidia
22-36 x 11-12 um, 4-spored, hyaline, yellow-brown or brown.
Pleurocystidia 34-50 x 6-8 um, fusiform with apical neck or
clavate-mucronate. Pseudocystidia 32-33 x 4-6 um, fusiform
and sometimes with apical neck or clavate, with golden
yellow pigmentation (plasmatic), along the tube edges;
occasionally a few pseudocystidia with size and shape of
the pleurocystidia, but with yellow pigment, on the tube
walls. All septa without clamps.
Cuticle a trichodermium formed of conical fascicles of
hyphae; thickness of cuticle 180 um; diameter of tricho-
dermial hyphae 3.5-5 um, hyphal walls deep red in Melzer's
reagent.
Stipe dermatocystidia variable, fusiform, with or
without a neck, or broadly clavate-mucronate; fusiform
cystidia 26-66(89) x (3)5-13 um; clavate-mucronate cystidia
30-35 x 14-18 wm; hyaline or golden yellow in KOH, brown in
Melzer's reagent. Tramal hyphae inamyloid.
Comments: see B. roseobadius
Collection examined:
Michigan, Washtenaw Co., Ypsilanti Golf Course,
Ypsilanti, in grassy oak woods, 3 Sep 1961, A.H. Smith
64046 - holotype, MICH.
DISCUSSION
There is not much variation in the spore size of the
different taxa studied here. However, the Q and Q values
are increasingly used as a taxonomic character (Breitenbach
and Kranzlin 1991, Wolfe 1991). B. pseudo-olivaceus with a
Q, of 2.4 differs from the other species with a Q, between
2eGoand) 258%
The hymenial cystidia range in shape from usually
fusiform to almost clavate. Fusiform cystidia often have an
apical neck. Following the terms used by Singer (1986) and
Wolfe (1991) for hymenial cystidia, most species have
fusiform or clavate, yellow pseudocystidia along the tube
edges and hyaline pleurocystidia located on the tube walls
and larger in size. Occasionally a few pseudocystidia can
be found on the tube wall, and they differ from the pleuro-
cystidia only in their yellow pigment. In contrast to the
genus Tylopilus Karsten, where hymenial cystidia play an
375
Figs. 16-20 Boletus subluridellus. 16 Spores. 17 Basidia.
18 Pseudocystidia. 19 Pleurocystidia. 20 Stipe dermato-
cystidia. Figs. 21-25 Boletus (TR 1316). 21 Spores.
22 Basidia. 23 Pleurocystidia. 24 Pseudocystidia.
25 Stipe dermatocystidia. Scale bars = 10m.
376
important role for the delimitation of species (e.g. Wolfe
1986, Wolfe 1991), the cystidia in the section Luridi do
not seem to be important taxonomic characters.
Stipe dermatocystidia have been used by Smith and
Thiers (1971) to delimit species in the section Luridi (B.
subluridellus, B. roseobadius, and B. rufocinnamomeus with
long, often setiform caulocystidia). Measurements taken
here show however, that they can range widely within one
collection and are often difficult to find, especially in
small collections. Some sporocarps of collection TR 1316
had very long stipe dermatocystidia but these were scat-
tered, difficult to detect and only present in some
fruitbodies of the collection. The shape of the stipe
dermatocystidia ranges from clavate to rostrate to fusiform
and their contents often stain brown in Melzer's reagent.
Basidia do not show variation between the species.
They are usually hyaline, but sometimes also pigmented.
Their basal septum is simple, without clamps.
The hymenophoral trama in all the species studied is
divergent, sometimes only slightly so, with a central
strand of parallel hyphae which stains brownish in Melzer's
reagent.
The cuticle can range from a trichoderm to a cutis.
Larger collections (TR 1316) show that there is variation
within species. The content of cuticle hyphae of B.
rufocinnamomeus turns brownish in Melzer's reagent. The
same character could occasionally be observed in the
colrectiuon. TRI 2a
In most cases the tips of trichodermial hyphae are
glued together to form conical fascicles.
A very conspicuous character of both B. subluridellus
and B. roseobadius are the cuticular hyphae many of which
have deep red walls in Melzer's reagent. However, this
constitutes no dextrinoidity, because the pigment can be
observed in water mounts as well. It can not be detected in
KOH, where it apparently changes its color or dissolves.
Rarely those pigmented hyphae could also be detected in the
cuticle of B. rufocinnamomeus. The pigment is often
indicated macroscopically by a red color of parts of the
fruitbody (e.g. pileus, stipe trama).
A peculiar character of B. pseudo-olivaceus is the
amyloid trama of stipe and pileus. Although very distinc-
tive, the intensity of the staining is not very strong and
quite homogeneous throughout the trama. A stronger amyloid
reaction in the genus Boletus can be found in cross walls
of tramal hyphae of Boletus calopus Fr. (Miller and Watling
1968) and in B. piedmontensis Grand and Smith in the form
of amyloid particles in the cuticular hyphae (Grand and
Smith 1971). .According “to “Smith and =Thiers, (1971)'«B.
pseudo-olivaceus is closely related to B. queletii Schulz.
from Europe, a view supported by Singer's (1977) observa-
tion of distinctly amyloid hyphae in the stipe of B.
queletii.
None of the species shows clamps in any parts of the
O77,
fruitbodies.
Themcollectson pik i316, wwhich has’ been included in
this study, seems to be close to or identical with B.
subluridellus. The presence of many red-walled hyphae in
the cutis and the spore measurements indicate a high
Similarity between the material from Virginia and collec-
tions of B. subluridellus. At the same time however, many
characters, like the pigmentation of the cuticular hyphae,
seem to vary even within collections of one species and may
depend on the age of sporocarps. Larger collections may
reveal the variability of some of the characters which are
traditionally used. Therefore a further study of a larger
number of other taxa in this group is obligatory and would
necessarily include the European taxa of this group.
ACKNOWLEDGEMENTS
I wish to thank Dr. C.B. Wolfe, Pennsylvania State
University, for helpful comments and a critical revision of
SHewmanuscCh pl uiMmeIndebGeneto Dre O.KeeMiller, 0%. for
hasenospitality ands lab! space. =f thank Dr. R.L. Shaffer,
curator of the Herbarium of the University of Michigan, for
providing loans. The "Deutsche Forschungsgemeinschaft"
(DFG) is acknowledged for providing a research grant.
LITERATURE
Breitenbach, J. and F. Kranzlin. 1991. Fungi of Switzer-
land. Vol. 3. Boletes and agarics. lst part. Edition
Mykologia, Lucerne, Switzerland.
Grand, L.F. and A.H. Smith. 1971. A new species of Boletus,
section Luridi, from North Carolina. Mycologia 63:
884-888
Kornerup, A. and J.H. Wanscher. 1967. Methuen Handbook of
Colour. Methuen and Co. Ltd., London, Great Britain.
Mebier, O-K., Jr.)and oR. | Watlinga FPl968.. The status . of
Boletus calopus Fr. in North America. Notes from the
Royal Botanic Garden Edinburgh 28: 317-325.
Singer, R. 1977. Keys for the identification of the species
of Agaricales I. Sydowia 30: 192-279.
Singer, R. 1986. The Agaricales in modern taxonomy. 4th ed.
Koeltz Scientific Books, Koenigstein, Germany.
Smith, A.H. and H.D. Thiers. 1971. The boletes of Michigan.
The University of Michigan Press, Ann Arbor, Michigan.
WOPlepeC=b. wi Loloashype studies tin Tylopilus-vltl Taxa
described by Walter H. Snell, Esther A. Dick, and co-
workers. Mycologias/s 2: 22-317
Wolfe, C.B., Jr. 1991. Type studies in Tylopilus (Boleta-
ceae). V. Taxa described by Alexander H. Smith, Harry
D. Thiers and Samuel J. Mazzer. Can. J. Bot. 69: 1833-
LOS. Or
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he 22
MYCOTAXON
Volume XLVI, pp. 379-385 April-June 1993
TWO NEW CRINIPELLINAE (TRICHOLOMATACEAE:
MARASMIAE) FROM SOUTH AMERICA
Roy E. HALLING
Institute of Systematic Botany
The New York Botanical Garden
Bronx, New York 10458-5126
ABSTRACT
Crinipellis ticoi from lowland Bolivia and Chaetocalathus magnus
from Andean Colombia are newly described from the Neotropics.
Both taxa possess dextrinoid, long, thick-walled, setoid hairs on the
pileus that distinguish these lamellate genera in the Crinipellinae.
Key Words: Agaricales, Crinipellinae, Crinipellis, Chaetocalathus,
Bolivia, Colombia
Study of specimens gathered in South America over a four year period (1986-
1990) has resulted in the discovery of many new taxa of agarics and boletes (eg.,
Halling & Ovrebo, 1987; Halling 1989a, 1989b, Singer et al., 1990; Horak & Hall-
ing, 1991; Tulloss et al., 1992; Baroni & Halling, 1992; Halling, 1992). Further ob-
servations on additional material has yielded a previously unknown species of
Crinipellis with bright orange pigmentation and the largest species of Chaetocala-
thus known to date. Fifty years ago, Singer (1942) provided a worldwide mono-
graph of these two genera, and more recently, published an updated monograph
for the Neotropical taxa (Singer, 1976). Two additional Neotropical species (one
from each genus) have since been recognized (Singer, 1989).
Color designations in parentheses (e.g., SA6) are taken from Kornerup and
Wanscher (1983). Other color terms (e.g. brick red) are general approximations of
those designations. Letter abbreviations describing spore sizes are: n = number of
spores measured, x = mean length x mean width, Q = mean length/width ratio.
Crinipellis ticoi Halling, sp. nov. Figs. 1-4
TYPE: BOLIVIA. Dpto. Beni: Prov. Iturralde, S of Rurrenabaque, Rio Tuichi near junction
with Rio Beni, “Laguna del Tigre,” +67°30’W, 14°25’S, 14 Apr 1990, Hailing 6433 (Holotype: LPB;
Isotype: NY).
Pileus aurantius sulcatus tomentosus vel subtomentosus, epilosus. Lamellae adnexae subdistantes
crassae latae aurantiae. Stipes centralis strictus +aequalis tenax tortilis pubescens vel fibrillosus vel
380
fibrilloso-pruinosus, pubescentis aurantiis, superfacies flava. Basidiosporae in cumulo albae, 12.1-14.3
x 5-7.1 um, inamyloideae. Hymenium pigmento aurantio in solutionibus alkalicis dissolventibus.
Pleurocystidia nulla. Cheilocystidia dispersa, hyalina, parietibus tenuibus et glabris. Setae pileipellis
sparsae dextrinoideae, parietibus crassis, interdum 2-3 septatae, apicibus obtusis vel subacutis vel sub-
capitatis. Setae stipitipellis abundantes dextrinoideae, parietibus tenuibus vel crassis, apicibus obtusis,
aliter setae pileipellis similis. Fibulae praesentes.
Etymology: In honor of Sr. Abellardo “Tico” Tudela R., of Rurrenabaque,
whose logistical support aided in the discovery of this species.
Pileus 1—3(-—4) cm broad, convex to plane, dry or moist and greasy then fading
when drying but not truly hygrophanous, orange (7A8—7B8) fading to a paler
orange (6A7-6B7 to 6A8-6B8 to near 5A6), sulcate to subsulcate to the disc,
even on disc, tomentose to matted subtomentose on disc, faintly so toward margin,
more yellowish on ridges with age and remaining orange in sulcae; margin even.
Flesh pale orange (5A2) in pileus, pale yellowish in stipe apex and below, up to 2
mm thick. Odor mild. Taste styptic but not unpleasant. Lamellae adnexed to sub-
free, subdistant, thick and broad (up to 7 mm), orange (7A8 to 6A8) but slightly
paler toward the even, nonmarginate edges; with lamellulae in 1-2 tiers. Stipe
(1—)2.5-7 cm long, (1—)3-—4 mm broad, insititious, strict, central, twisted, tough,
+equal but often broadest at apex and base, sometimes with a bulb at base; sur-
face pubescent to fibrillose to fibrillose-pruinose overall, denser toward the base,
with orange (color range as cited above) fibrils on yellowish ground above and be-
low when young, brown toward base with age, sometimes cream-colored slightly at
base; interior stuffed, not hollow.
Fig. 1. Crinipellis ticoi. (Halling 6393). x1.
381
5 um
Figs. 2-4. Microscopic features of Crinipellis ticoi (Halling 6433). 2. Basidiospores. 3.
Cheilocystidia. 4. Setae from pileipellis.
Basidiospores white in deposit, (10—)12.1—14.3 x 5—7.1 wm (n = 20, x = 13 x
5.8 um, Q = 2.3), ellipsoid to suballantoid and inequilateral in profile, ellipsoid to
subcylindric in abaxial view, inamyloid and acyanophilous, smooth, thin-walled.
Hymenium embedded in an amorphous golden orange pigment that dissolves in
alkali. Basidia 28-35 x 7-8.4 um, clavate, 4-sterigmate, not siderophilous.
Pleurocystidia absent. Cheilocystidia scattered, inconspicuous, hyaline, thin-walled,
smooth, subfusoid to subcylindric, 38-49 x 6-8 wm. Lamellar trama interwoven, |
inamyloid; hyphae smooth, thin-walled, 3.5-12.6 um diam. Pileus trama radially
oriented, inamyloid; hyphae smooth, thin-walled, S—10(-14) um diam. Setae
sparsely distributed on pileus surface especially toward the margin, more abundant
on the disc, 70-210 x 5—9 um, thick-walled, dextrinoid, occasionally 2—3 septate,
with obtuse to subacute to subcapitate apices, arising from a pileipellis of radially
oriented, repent, thin-walled, hyaline hyphae; cells 7—14(-17.5) um in diam,
sometimes lightly encrusted and slightly inflated. Setae abundant on stipe surface,
Strict or curved, dextrinoid, thin- to thick-walled, with obtuse apices, otherwise
resembling setae in pileipellis, arising from a stipitipellis whose elements are verti-
cally oriented, dextrinoid, smooth, and thin-walled, 3.5-—6 um diam. Clamp con-
nections present.
On rotten wood in lowland Amazonian rainforest.
382
Additional material examined: BOLIVIA. Dpto. Beni: Prov. Iturralde, S of Rurrenabaque, along
Rio Beni between town of Rurrenabaque and junction of Rio Tuichi, +67°30’W, 14°25’S, 4 Apr 1990,
Halling 6393 (LPB, NY). Prov. Gral. José Ballivian, +1 km E of Rurrenabaque, along road to air-
port, +67°30’W, 14°20’S, 29 Mar 1990, Halling 6372 (LPB, NY); S of Rurrenabaque, along Rio Beni
between town of Rurrenabaque and junction of Rio Tuichi, +67°30’W, 14°25’S, 3 Apr 1990, Halling
6383 (LPB, NY), 7 Apr 1990, Halling 6416 (LPB, NY).
Crinipellis ticoi seems best accommodated in sect. Jopodinae (Singer, 1976)
characterized by the brightly pigmented pileus and tissues not greening or graying
in alkali. In this section, C. insignis Sing., C. purpurea Sing., and C. hygrocybioides
(Henn.) Sing. appear most similar. Crinipellis insignis differs in the purple to pur-
plish pink pileus, white lamellae, and short, eccentric stipe. Crinipellis purpurea is
distinguished by the dark purple red pileus, white lamellae, and smaller spores
(6.5-10.3 x 3-4.7 um). Also, both of these species are described as having
abundant setae giving the pileus a pilose aspect. Crinipellis ticoi has a tomentose
pileus with scattered setae. Comparison of C. ticoi to C. hygrocybioides is more
problematical. When Singer (1989) transferred Marasmius hygrocybioides Henn. to
Crinipellis, he did not provide further data or documentation other than to indicate
that the type is a species of sect. Ionopodinae [sic]. Henning’s (1902) original des-
cription tersely describes a yellow orange, small, marasmioid fungus. Two isotypes
(Kamerun. Bipindi, Mai 1899, G. Zenker 2039, PC!; K!) have been compared. The
Kew material consists of one basidiome which was not examined microscopically.
The Paris specimen contains some 14 complete basidomata and additional stipes
lacking pilei. Abundant dextrinoid setae are present on the pileus and stipe in-
dicating Crinipellis, but spores and cystidia were not observed. Contrary to C. ticoi,
C. hygrocybioides is a smaller fungus (pileus 6-11 mm broad, stipe 10-20 x
0.5-0.8 mm) and has an umbilicate to papillate pileus which is pilose at the
margin.
Chaetocalathus magnus Halling, sp. nov. Figs. 5-8
TYPE: COLOMBIA. Dpto. Antioquia: Mpio. Guarne, near Centro Experimental Piedras
Blancas, 14 km E of Medellin, 75°29’W, 6°15’N, 2350 m, 28 Nov 1988, Halling 6159 (Holotype: HUA;
Isotype: NY).
Basidiomata estipitata. Pileus siccus pigmento brunneo, primo planus et minime granulosus vel
velutinus tum rugulosus vel subsulcatus. Lamellae primo confertae et albae tum subdistantes et
aurantio-albae. Basdiosporae 9.3-12.1 x 6.4-8.6 um, dextrinoideae et cyanophilae. Pleurocystidia
rarissima, inamyloidea, aliter cheilocystidia simili. Cheilocystidia inferme dextrinoidea, nonincrustata,
parietibus crassis vel tenuibus, appendiculis apicalibus. Setae pileipellis dextrinoideae, parietibus crassis,
apicibus obtusis. Fibulae praesentes.
Etymology: Latin magnus, great or large.
Basidiomata nonstipitate. Pileus (0.7-)1-—2.2(-—2.7) cm broad, pendant-
ungulate to barely laterally attached, dry, even and matted velutinous to minutely
granulose and burnt sienna to brick red (7D8-—7) when young, soon rugulose to
subsulcate and fading to near grayish orange (6BS5), white to tan near attachment
point. Flesh thin, pallid. Odor and taste mild. Lamellae close and white when
young with unequal widths and barely subfimbriate edges, soon subdistant and
orange white, afimbriate, subintervenose.
383
Fig. 5. Chaetocalathus magnus (Halling 6159). x2.
Figs. 6-8. Microscopic features of Chaetocalathus magnus (Halling 6159). 6. Basidiospores. 7.
Cheilocystidia. 8. Setae from pileipellis.
384
Basidiospores 9.3-12.1 x 6.4-8.6 um (n = 20,x = 10.4 x 7 um, Q = 1.5)
broadly ellipsoid, sometimes subamygdaliform, weakly dextrinoid when observed-
singly, more strongly so in mass, cyanophilous, hyaline in KOH, smooth, thin-
wailed, with a slightly thickened wall in age, occasionally transversely septate.
Basidia 28-35 x 8-9.5 um, clavate, hyaline, thin-walled, not siderophilous, 4-
sterigmate, Pleurocystidia inconspicuous and extremely rare, thick-walled, in-
amyloid, otherwise resembling cheilocystidia. Cheilocystidia 25-46 x 10-15 um,
thin-walled or thick-walled, with 2—4 apical appendages, typically subcylindric, not
encrusted, usually inamyloid or very rarely weakly dextrinoid, and then the reac-
tion extremely localized on a given cystidium. Lamellar and pileal trama inter-
woven, hyaline, inamyloid, not gelatinous; hyphae 5-7 um broad, filamentous,
thick-walled (up to 3 wm). Pileipellis an interwoven layer of repent, hyaline, in-
amyloid hyphae, 4—8 um in diam, lightly encrusted with pigment, thin-walled to
progressively thicker-walled as they give rise to setoid hairs; setae 100-200 um x
3.5-7 ym, thick-walled, sometimes entirely lacking a lumen, dextrinoid, smooth,
highly refractive to pale golden in KOH, with obtuse apices. Clamp connections
present.
On stems of Vaccinium (Ericaceae) in Quercus humboldtii forest.
Additional material examined: COLOMBIA. Dpto. Antioquia: Mpio. Guarne, near Centro Ex-
perimental Piedras Blancas, 14 km E of Medellin, 75°29’W, 6°15’N, 2350 m, 10 Nov 1988, Desjardin
4763 (HUA, SFSU).
This is the eighth species of Chaetocalathus known from the Neotropics, and
the second known from Colombia [after the widespread Ch. liliputianus (Mont.)
Sing.]. Chaetocalathus magnus fits best in sect. Holocystis Sing. by virtue of the
dextrinoid (rarely), thick-walled, apically appendaged cystidia. However, it can be
distinguished easily by its large size, brown pigmentation of the non-pilose pileus,
non-encrusted cystidia, and lack of entire and simple pleurocystidia. However,
when pleurocystidia are present, they are extremely rare and resemble the in-
amyloid cheilocystidia.
ACKNOWLEDGMENTS
Field work in Bolivia and Colombia was supported by National Science Foun-
dation Grant #BSR — 8600424. I am indebted to the staff and students at Univer-
sidad de Antioquia (HUA) for aid and guidance in Colombia; for assistance in
Bolivia, I thank Emilia Garcia (LPB), the late David Smith (MO), David and
Sandy Williams (USDA), and Sr. Abellardo “Tico” Tudela R. in Rurrenabaque.
Gregory Mueller (F) is thanked for his review of this paper. Further, I am grateful
to Dennis E. Desjardin (SFSU) and Ana E. Franco-M. (NY) who provided out-
standing field assistance.
LITERATURE CITED
Baroni, T. J. & R. E. Halling. 1992. New species of Rhodocybe from South America with a key to
species. Mycologia 84: 411-421.
385
Halling, R. E. 1989a. A synopsis of Colombian Boletes. Mycotaxon 34: 93-113.
1989b (1990). Notes on Collybia III. Three neotropical species of subg. Rhodocollybia.
Mycologia 81: 870-875.
. 1992. A new species of Boletus section Luridi from Colombia. Brittonia 44: 108-113.
, & C. L. Ovrebo. 1987. A new species of Rozites from oak forests of Colombia with notes on
biogeography. Mycologia 79: 674-678.
Hennings, P. 1902. Fungi camerunensis novi. III. Bot. Jahrb. Syst. 30: 39-57.
Horak, E. & R. E. Halling. 1991. New records of Phaeocollybia from Colombia. Mycologia 83: 464-
472.
Kornerup, A. & J. H. Wanscher. 1983. Methuen Handbook of Colour. Ed. 3. Eyre Methuen Ltd.,
London.
Singer, R. 1942. A monographic study of the genera Crinipellis and Chaetocalathus. Lilloa 8:
441-534.
. 1976. Marasmieae (Basidiomycetes —Tricholomataceae). Flora Neotrop. Monogr. 17: 1-347.
. 1989. New taxa and new combinations of Agaricales (Diagnoses fungorum novorum Agari-
calium IV). Fieldiana, Bot., n.s. 21: 1-133.
, C. L. Ovrebo, & R. E. Halling. 1990. A new species of Phylloporus and a new species of
Tricholomopsis from Colombia with notes on Phylloporus boletinoides. Mycologia 82: 452-459.
Tulloss, R. E., C. L. Ovrebo, & R. E. Halling. 1992. Studies on Amanita (Amanitaceae) from
Andean Colombia. Mem. New York Bot. Gard. 66: 1-46.
ra Sar.
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MY COTAXON
Volume XLVI, pp.387-388 April-June 1993
SEPTORIA THYMI SP. NOV. FROM BULGARIA
Ekaterina F. Sameva and Ganka G. Bakalova
Institute of Botany, 1113 Sofia, Bulgaria
ABSTRACT. A new species of the genus Septoria Sacc.
(Coelomycetes), Septoria thymi Sameva et Bakalova
sp. nov. from Bulgaria is described and illustrated.
During inventory investigations of the genus Septoria Sacc.
(Sphaeropsidales, Coelomycetes) in Bulgaria, a new species belonging
to this genus is established.
Septoria thymi Sameva et Bakalova, sp. nov.
Maculae 0.1-0.3 cm in diam, orbiculares vel irregularis,
solitariae, albidis, ochraceae vel pallide-brunneae indistincto
atropurpureo marginate. Pycnides epiphylae, globosae vel depresso-
globosae tenuiter tunicatae, pallido-brunneae, 140-180 um in diam,
poro orbiculari 10-20 um in diam, cellulis parvis obscurioribus cincto.
Conidia cyindrica, elongato-fusiformia, clavata, utrinque rotundata,
recta vel leniter curvata, 3-5(7) septata, hyalina, 30.0-(X =38.94;
S =4.31)-54.0 x 2.5-(X =3.41; S=0.43)-5.0 um, N=100 (Figs. 1, 2).
| In foliis Thymi pulegioidi L., Bulgaria, mons Vitosha, supra
pagum Bistrica, 13.07.1980, E. F. Sameva, G. G. Bakalova, SOM 20763
M, holotypus.
In speciebus Thymi Septoria primum observatur.
Leaf spots 0.1-0.3 cm in diam, rounded or irregular, single,
whitish, ochraceus or pale brown, surrounded by a brown or purple
border. Pycnidia epiphyllous, spheric or flattened, with thin walls, pale
brown, 140-180 um in diam, with circular ostiole, surrounded by darker
cells, 10-20 wm in diam. Conidia cylindric, fusiform or clavate,
rounded at both ends, straight or slightly curved, 3-5 (rarely 7) septate,
hyaline, 30.0-(X-38.94; S=4.31)-54.0 x 2.5-(X =3.41; $=0.43)-5.0 um.
N=100 (Figs. 1, 2).
388
On leaves of Thymus pulegioides L., Bulgaria, Mount Vitosha,
above the village of Bistritsa, 13.07.1980, E. F. Sameva, G. G.
Bakalova, SOM 20763 M (holotype).
First description of a species of Septoria Sacc. on Thymus L.
Figs. 1, 2. Pycnidium and conidia of Septoria Thymi
MY COTAXON
Volume XLVII, pp. 389-394 April-June 1993
THREE NEW SPECIES AND A NEW VARIETY OF
PLUTEUS FROM THE UNITED STATES
PARTHA BANERJEE! AND WALTER J. SUNDBERG
Department of Plant Biology, Southern Illinois University,
Carbondale, Illinois 62901, U.S.A.
‘Present Address: Biological Survey, Cultural Education Center,
New York State Museum, Albany, NY 12230, U.S.A.
ABSTRACT: Three previously unreported taxa, Pluteus cinerascens, P. heterocystis,
P. laricinus, and P. salicinus var. americanus were discovered during field and herbarium
studies of Pluteus section Pluteus (Pluteaceae, Agaricales) in the United States. They are
described below as new.
KEY WORDS: Agaricales, Pluteus, United States.
The genus Pluteus Fr. (Pluteaceae, Agaricales) is a large group of presumably non-
mycorrhizal mushrooms found on all continents excepting Antarctica (Singer 1986). The
genus has three sections: Pluteus Fr., Hispidoderma Fayod and Celluloderma Fayod. In
section Pluteus, the pileipellis is a filamentous cutis with cylindric or cylindric-fusoid end-
cells; the hyphae are appressed with the end-cells sometimes erect or suberect in fascicles.
The pleurocystidia are metuloid. Metuloid cystidia are thick-walled, veniricose (i. e.,
swollen in the middle), and usually with apical projections.
In the following descriptions, color names within quotes are from Ridgway (1912,
R); Kelly and Judd (1976, = NBS); and the Royal Botanic Garden Color Chart (1969,
RBG). Color names without quotes are author-generated.
All anatomical observations were made in 3% KOH.
The following are definitions used to describe pleurocystidia: "cervinus-type" refers to
pleurocystidia with apical ornamentations while "magnus-type" indicates pleurocystidia
without apical ornamentations (modified after Singer 1956).
Unless otherwise noted, herbarium abbreviations (Holmgren et al. 1990) for specimens
examined are given in parentheses after the specimen number. Standard two-lettered
abbreviations are used for American states.
390
Pluteus cinerascens P. Banerjee et W. J. Sundberg, sp. nov. Fig. 1.
Speciebus aliis sectionis Plutei, pileo 2-7 cm lato, margine cinerasci ubi vulnerato
(injurio vel contuso); cervini-pleurocystidiis metuloideis ventricosis, tunica #2.2 Lm
crassa, apice acuto vel truncato, appendiculo, cornibus 2-4 plerumque fortibus longis
acutis; epicutis pilei fibulis destituta.
Holotypus: A. H. Smith 57433! (MICH). Solitary on a very rotten log, Proud Lake
Recreation Area, Oakland Co., Michigan, U.S.A.
Pileus 2-7 cm broad, broadly convex with a slightly flattened disc, surface with small
fascicles of appressed, cinnamon-buff fibrils on a white ground color over the disc, white
and glabrous elsewhere, a dingy orange flush developing in some areas overnight, injured
and bruised places along margin becoming "cinereous" (R) to "clay buff" (RBG); context
white, soft; odor and taste raphanoid. Lamellae free and remote from stipe, close, broad,
white becoming dingy pale "vinaceous" (R) to "vinaceous buff" (RBG), edges serrulate,
concolorous with lamellae, beaded with drops when fresh; lamellulae 1-2-tiered. Stipe 6.5-
10 cm long, 0.2-1 cm thick at apex, base cylindrical to clavate and up to 1.5 cm broad,
naked above, lower third may or may not have appressed fibrils, fibrils white overall but
staining or discoloring gray to fuliginous in injured places and with dingy "ochraceous" (R)
streaks over the base; solid but fragile.
Basidiospores 6-7.4 x 4.4-5.6 jim, oval-ellipsoid, wail smooth, moderately thick,
rarely collapsing, with hyaline cytoplasmic content, some with a large lipid droplet; hilar
appendix inconspicuous. Basidia 20-28 x 7-8.4 um, clavate; four-spored. Pleurocystidia
very abundant, (30-)42-85 x 12-22 um, ventricose, thick-walled (1.4-2.0 um), almost all
hyaline ("content pale lilac in KOH when fresh", Smith, unpublished); of cervinus-type,
apex acute to truncate and with (1-)2-4(-5) well-developed, mostly long and sharp apical
hooks, some pleurocystidia with short to blunt apical hooks, lateral spinules occasionally
present, occasional sphaeropedunculate pleurocystidia interspersed with those of the
cervinus-type; pleurocystidia near lamellar edge mostly of cervinus-type interspersed with a
few magnus-type. Cheilocystidia not abundant to abundant, in fascicles when abundant;
30-65 x 12-22 um, sphaeropedunculate, hyaline; with a tendency to gelatinize when
remoistened. Pileipellis hyphae 4-12 tm in diameter, ends rounded, most hyaline, some
with a homogeneous dark to light brown content; pellicle in cross section easily separates
from the context; clamp connections absent on pileipellis hyphae.
Other collections examined: Klavins s. n., Pomona Natural Bridge, Jackson Co., IL
(SIU).
Observations: The morphological and anatomical characters are significant enough to
describe it as a new species. A. H. Smith (unpublished) considered P. cinerascens close to
P. magnus McClatchie, but the presence of very strongly developed long, sharp
pleurocystidial hooks indicate closer affinities to P. cervinus (Schaeff.: Fr.) Kumm. The
cinerous Stains and the strongly developed hooks on many of the pleurocystidia distinguish
it. A solitary liginicolous basidiocarp of the type specimen was collected from Michigan by
A. H. Smith, and another solitary terrestrial basidiocarp has been found recently in Illinois.
Figs. 1-4. Anatomical characters of Pluteus species. 1. Pluteus cinerascens sp. nov.; 2.
P. heterocystis sp. nov.; 3. P. laricinus sp. nov.; 4. P. salicinus var. americanus var. nov.
(P: Pleurocystidia; C: Cheilocystidia; L: Pileipellis; B: Basidia; S: Basidiospores.
Scale lines: 10 um for basidiospores; 20 um for other structures).
392
The morphological description as well as notes on its habit, habitat and distribution of
the type specimen are adapted from data obtained by A. H. Smith (unpublished).
Pluteus heterocystis P. Banerjee et W. J. Sundberg, sp. nov. Fig. 2.
Speciebus aliis sectionis Plutei, pileo 5 cm lato, disco umbrino et fusco fibrillulo in
Strias versus marginem radiatim; lamellis crassis marginibus cariosis; pleurocyStidiis
metuloideis clavatis late ventricosis, apice obtuso vel rotundato, appendiculo cornibus
brevibus vel nullis; epicutis pilei fibulis destituta.
Holotypus: A. H. Smith 13616! (MICH). Solitary on soil, Hoh River, Olympic
National Park, Washington, U.S.A. Known only from the type collection.
Pileus 5 cm broad, expanded-umbonate, umbo obtuse, moist to subviscid (as in
Inocybe geophylla); disc "Saccardo's umber" (R) and with dark fibrils extending as streaks
toward the margin; marginal area watery gray but opaque; odor and taste not recorded.
Lamellae free, remote from the stipe, close, moderately broad, somewhat thick, white
becoming dingy pink; edges eroded and pallid; lamellulae 1-2-tiered. Stipe 7 cm long, 1
cm thick at apex, equal to a slightly bulbous base, glabrous over most of the surface, some
"umber" (RBG) fibrillose streaks present on lower portion; solid; context pallid.
Basidiospores 6.6-8(-9) x 5.6-6.4(-7) um, broadly ovate to ellipsoid, smooth, wall
slightly thick, some hyaline and with a large lipid droplet, some with cytoplasmic content;
hilar appendix visible. Basidia 24-32 x 8-10 um, clavate, usually 4-spored, occasionally
2-spored, hyaline. Pleurocystidia abundant and dimorphic: one form 50-80 x 10-25 um,
clavate to broadly fusoid-ventricose, apices more or less truncate, many with several short
horns at apex and others merely with aborted apical ornamentation; the second form with
apex rounded and lacking horns, wall uniformly 1.2-1.8 um thick, no lateral ornamentation
seen; all pleurocystidia hyaline., Cheilocystidia 35-45(-54) x 8-12.6 um, clavate-pedicellate
to slightly ventricose, thin-walled, hyaline; apices obtuse to rounded, smooth. Pileipellis
of radial hyphae 6-10 tm in diameter; end cells often subfusoid and 9-14 tm in diameter;
content hyaline or rarely dingy yellow-brown, rarely a few hyaline pilocystidia (15-25 um
diameter) visible; clamp connections not seen on pileipellis hyphae.
Observations: The thick gills with eroded edges, and the presence of two types of
pleurocystidia with one form having truncated apices with short ornamentations are
distinctive. A. H. Smith (unpublished) found clamp connections in the pileipellis, but our
repeated attempts failed to demonstrate them.
The morphological description as well as data on its habit, habitat and distribution are
modified from notes of A. H. Smith (unpublished).
Pluteus laricinus P. Banerjee et W. J. Sundberg, sp. nov. Fig. 3.
Speciebus aliis sectionis Plutei, pileo 1.5 cm lato, fusci-umbrino, squamuloso in disco;
lamellis albis ad marginem pilei griseolis; epicute pilei fibulis infrequentibus; stipite gracili,
0.2 cm in diametro ad apicem.
Holotypus: A. H. Smith 38888! (MICH). Solitary on a rotten larch log. Proud Lake
Recreation Area, Oakland Co., Michigan, U.S.A. Known only from the type collection.
393
Pileus 1.5 cm broad, convex-umbonate, squamulose on the disc, elsewhere appressed-
fibrillose, dark "wood brown" (R); margin slightly wavy but not striate. Lamellae broad,
remotely free, close, white, becoming pale pink and edges grayish near cap margin;
lamellulae 1-2-tiered. Stipe 3 cm long, 2 cm thick at apex, only slightly enlarged
downward, whitish and not darkening, with faint "avellaneous" (R) pruina near base.
Basidiospores (4.4-)5-6.4 x 4.6-5 1m, oval-ellipsoid, smooth, wall only slightly thick,
hyaline or nearly so; hilar appendix visible. Basidia 22-25 x 6-7.4 tm, clavate, four-
spored, hyaline. Pleurocystidia 35-75 x 8-20 tm, fusoid-ventricose, hyaline, wall of
upper part thicker (1.2-1.5 jum thick) than lower part; most pleurocystidia near the gill edge
are of magnus-type (i.e., without hooks), others with 2-3 hooks at the apex, rarely with 1
lateral hook. Cheilocystidia 28-48 x 10-14.6 um, clavate-pedicellate to broadly fusoid,
smooth and apices rounded, walls thin, content hyaline to dingy brownish. Pileipellis
hyphae 3-5 tm in diameter, with rounded to subacute apices, occasionally hyphal end cells
fusoid-ventricose. Pilocystidia 85-150 x 8-14 um, deeply embedded in the pileal trama,
content of all pileipellis hyphae hyaline or dull yellow brown; clamp connections on
pileipellis hyphae present but not frequent.
Observations: The small spores, the grayish gill margin, and slender stipe distinguish
the species. A. H. Smith (unpublished) found abundant clamp connections on pileipellis
hyphae, but after repeated attempts, we found only a few.
The morphological description as well as notes on its habit, habitat and distribution are
adapted from notes by A. H. Smith (unpublished).
Pluteus salicinus var. americanus P. Banerjee et W. J. Sundberg, var. nov. Fig. 4.
Varietatibus aliis Plutei salicini, pileo 5-6 cm lato, glabro hygrophano, ad marginem
translucidi-striato; pleurocystidiis acutis vel truncatis, appendiculis cornibus apicis
complexis; epicute pilei fibuligera.
Holotypus: A. H. Smith 57841! (MICH). Solitary on aspen wood. Pellston Hills,
west of Pellston, Michigan, U.S.A.
Pileus 5-6 cm broad, broadly convex expanding to plane with no umbo, color between
"drab" (R) and "hair brown" (R), surface moist and hygrophanous, sometimes turning
greenish-gray upon handling or drying, opaque when faded; lacking any spinules or
squamules over disc; margin translucent-striate fading to an ashy gray; context watery
grayish fading to pallid; odor of crushed context disagreeable, taste very disagreeable:
somewhat metallic. Lamellae fairly broad, remote, close, whitish pallid, "pink" (NBS) in
age; edges concolorous; lamellulae 1-2-tiered. Stipe 4-6 cm long, 0.5-0.6 cm thick, silvery
whitish to grayish and base soon staining greenish gray when handled or dried; naked;
hollow, terete or compressed.
Basidiospores 8-10(-11) x 6.4-7.4 um, ellipsoid to ovoid, smooth, hyaline, wall
slightly thick; hilar appendix conspicuous. Basidia 28-34 x 7-8.8 tm, clavate, both 2- and
four-spored. Pleurocystidia abundant, 50-85 x 12-28 1m, ventricose to very broadly
fusoid with apex more or less truncate, some with simple sharp and pointed projections,
some others with short (2-5 zm) truncate projections arising around the neck and these
usually with a crown of 4-5 small but pronounced projections around their apices (i.e.,
primary and secondary ornamentations present), rarely some aborted ornamentations
observed on the neck just below the apex; wall from ventricose portion downward to
pedicel becoming thin and readily collapsing, 1.2-1.4 um thick near neck, +1 um thick
near pedicel; all pleurocystidia hyaline. Cheilocystidia in fascicles, 34-46 x 12-20 um,
394
sphaeropedunculate to clavate-pedicellate, thin-walled, smooth, hyaline. Pileipellis a
trichoderm formed of dimitic hyphae 3-6 1m in diameter, occasionally with cystidioid end-
cells up to 16 tm in diameter, content pale brownish "fuscous" (R) to dingy brown; clamp
connections present on pileipellis hyphae.
Other collections examined: A. H. Smith 57846, MI (MICH); A. H. Smith 57099, MI
(MICH); A. H. Smith 57787, MI (MICH)—no other data available on these collections;
TN 14277, Indian Creek woods, Great Smoky Mountains National Park, Swain Co., NC
(TENN).
Observations: This taxon is in the Pluteus salicinus (Pers.: Fr.) Kumm. group, but is
distinctive because of its hygrophanous pileus with translucent-striate margin when moist.
Also, the presence of some pleurocystidia with compound apical ornamentations is
distinctive. Among the other recognized American varieties of P. salicinus, P. salicinus
var. salicinus is not hygrophanous, and P. salicinus var. achloes Sing. is occasionally
hygrophanous. Moreover, these two varieties almost never have pleurocystidia with
compound apical ornamentations. InP. salicinus var. americanus, the stipe (and
sometimes pileus) turn dark bluish-fuscous upon handling or drying which is quite
distinctive for a Pluteus but this character is present to some extent throughout the P.
salicinus group.
ACKNOWLEDGEMENTS
We wish to thank Dr. Gregory M. Mueller of the Field Museum of Natural History for
his pre-submission review of the manuscript. We express our gratitude to Dr. Charles
Sheviak and Ms. Patricia Kernan of the New York State Museum Biological Survey for
their help with the latin diagnoses and illustrations, respectively. The University of
Michigan Herbarium is thanked for the loan of the Pluteus specimens.
LITERATURE CITED
Henderson, D. M., P. D. Orton, and R. Watling. 1969. Colour Identification Chart.
British Fungus Flora. Agarics and Boleti: Introduction. Royal Botanic Garden,
Edinburgh. 84 pl.
Holmgren, P. K., N. H. Holmgren, and L. C. Barnett (Eds.). 1990. Index
Herbariorum. Part I. 8th Ed. New York Botanical Garden, Bronx. 693 p.
Kelly, K. L., and D. B. Judd. 1976. Color. Universal Language and Dictionary of
Names. Nat. Bur. Stand. Spec. Publ. 440. U.S. Gov't. Printing Office, —
Washington, D.C. 158 p.
Ridgway, R. 1912. Color Standards and Color Nomenclature. Publ. by the author.
Washington, D.C. 43 p.+ 53 pl.
Singer, R. 1956. Contributions toward a monograph of the genus Pluteus. Trans. Brit.
Mycol. Soc. 39(2): 145-232.
Singer, R. 1986. The Agaricales in Modern Taxonomy. 4th ed. Koeltz Scientific Books,
Koenigstein, F.R.G. 981 pp.
MY COTAXON
Volume XLVI, pp. 395-403 April-June 1993
WORLD LIST OF CETRARIOID LICHENS
TIINA RANDLANE & ANDRES SAAG
Institute of Botany and Ecology, Tartu University
Lai Street 38, Tartu, Estonia EE-2400
Abstract. The list sums up 120 cetrarioid lichen species. The
present generic location and valid epithet is indicated for each of
them. The generic location of 27 species (marked with *) is not
acceptable but not yet converted. Information available about 9
species (marked with ?) is unsufficient.
This is the first attempt after 40 years (Rassadina, 1950; Rdsdnen, 1952) to
compile a total list of cetrarioid lichens. The terms "cetrarioid", "parmelioid" and
"alectoroid lichens" are nowadays in wide use but not strictly defined. All those
terms indicate the certain morphological trends in the fam. Parmeliaceae,
"cetrarioid" being perhaps the most unlimited of the three. Classically foliose thallus
with dorsiventral structure which is not closely adnate to the substrate is
characteristic to this trend as well as dominating marginal position of apothecia and
pycnidia. We have followed this classical treatment and comprised into the list of
cetrarioid lichens all the species corresponding to those general characters.
According to the contemporary taxonomy the cetrarioid lichen species may belong
to different genera while most of them have earlier been included in the genus
Cetraria. The latter was described in 1803 by Erik Acharius and consisted then of
8 species. The number of species in the genus has varied greatly being the biggest
(76) in the treatment of the Russian lichenologist Ksenya Rassadina in 1950. At
present the obvious trend in the systematics of the fam. Parmeliaceae is towards the
delimitation of small and more or less homogeneous genera. During last decades
13 separate genera of cetrarioid lichens have been newly described or resurrected:
Ahtiana (Goward, 1985), Allocetraria (Kurokawa & Lai, 1991), Asahinea (Culberson
& Culberson, C.,1965), Cetrariopsis (Kurokawa, 1980), Cetrelia (Culberson &
Culberson, C., 1968), Cetreliopsis (Lai, 1980), Esslingeriana (Hale & Lai in Lai,
1980), Masonhalea (Karnefelt, 1977), Nephromopsis (Miller Argoviensis, 1891;
resurrected by Lai in 1980), Parmelaria (Awasthi, 1987), Platismatia (Culberson &
Culberson, C., 1968), Tuckermannopsis (Gyelnik, 1933; resurrected by Hale in Egan,
1987) and Vulpicida (Mattsson & Lai, 1993). Among these units the genera Cetraria
396
and Tuckermannopsis are still the conglomerates of poorly connected taxa and in
urgent need of further studies. Description of some more new cetrarioid genera are
expected in the near future.
In our list the specific epithets are presented in the alphabetical order. The
present generic location and valid epithet is indicated for each of them. All together
120 species are accepted as separate. Their epithets are numbered. The epithets
without numbers are treated as synonyms, or they represent the species of other,
not cetrarioid genera. There is not much information available about 9 species
which may therefore turn out to be either synonyms or belong even to quite
different genera. These species are marked with ? in the list. The species which
present generic location is not acceptable but not yet converted, are marked with
* The synonymity of some species is still doubtful. On these occasions the marks
=? are used.
aculeata (Schreber) Fr. [Cetraria] = Coelocaulon aculeatum (Schreber) Link
*1. agnata (Nyl.) Krist. [Cetraria] = Cetraria agnata (Nyl.) Krist.
2. alaskana Culb.& C.Culb. [Cetraria] = Cetrelia alaskana (Culb. & C.Culb.)
Culb.& C.Culb.
?3. albopunctata Zahlbr. {Cetraria]
aleurites (Ach.) Th.Fr. [Cetraria] = Imshaugia aleurites (Ach.) S.F.Meyer
alvarensis (Wahlenb.) Lynge [Cetraria] = Vulpicida tubulosus (Schaerer)
J.-E.Mattsson & Lai
4. ambigua Bab. [Cetraria] = Allocetraria ambigua (Bab.) Kurok. & Lai
americana Sato [Cetraria| = Tuckermannopsis orbata (Nyl.) Lai
5. americana Sprengel [Nephroma] = Tuckermannopsis americana (Sprengel)
Hale
6. andrejevii Oxner [Cetraria] = Cetraria andrejevii Oxner
?7. annae Oxner [Cetraria]
28. antarctica Zahlbr.
aradensis Gyelnik [Pseudoparmelia| = Cetrelia olivetorum (Nyl.) Culb. &
C.Culb.
arborialis (G.K.Merr.) Howard [Cetraria] = Cetraria subalpina Imsh.
arctica (Hook.) Tuck. [Cetraria] = Dactylina arctica (Hook.) Nyl.
arctica Magnusson [Cetraria] = Cetraria nigricascens (Nyl.) Elenkin
9. arenaria Karnef. [Cetraria] = Cetraria arenaria Karnef.
10. asahinae Sato [Cetraria] = Nephromopsis asahinae (Sato) Raésdénen
atlantica (Tuck.) Du Rietz [Cetraria] = Platismatia tuckermanii (Oakes) Culb.
& C.Culb.
*11. aurescens Tuck. [Cetraria] = Tuckermannopsis aurescens (Tuck.) Hale
12. australiensis W.Weber ex Karnef. [Cetraria] = Cetraria australiensis W.Weber
ex Karnef.
bavarica Krempelh. [Cetraria] = Tuckermannopsis oakesiana (Tuck.) Hale
billardieri Mont. [Cetraria] = Hypogymnia billardieri (Mont.) Filson
bohemica Anders [Cetraria] = Coelocaulon aculeatum (Schreber) Link
13. braunsiana (Mill. Arg.) Zahlbr. [Cetraria] = Cetrelia braunsiana (Miill. Arg.)
Culb. & C.Culb.
californica G.K.Merr. [Cetraria] = Tuckermannopsis merrillii (Du Rietz) Hale
*14,
15.
16.
1;
*18.
Lo
20.
ZA
B22.
Pao
24.
2).
397
californica Tuck. [Cetraria] = Cetraria californica Tuck.
californica Gyelnik [Nephromopsis| = according to the description (Gyelnik,
1931) ought to be Tuckermannopsis orbata (Nyl.) Lai. This is confirmed
by TLC of isotype (FH) which contains protolichesterinic acid.
canadensis (Rasanen) Rasanen [Cetraria] = Vulpicida canadensis (Rasaénen)
J.-E.Mattsson & Lai
caperata sensu Vainio [Cetraria] = Vulpicida pinastri (Scop.) J.-E.Mattsson
& Lai
capitata Lynge [Cetraria] = Cetraria nigricans Nyl.
cavernosus Menzies [Lichen] = Platismatia lacunosa (Ach.) Culb. & C.Culb.
cetrarioides (Delise ex Duby) Nyl. [Parmelia] = Cetrelia cetrarioides (Delise
ex Duby) Culb. & C.Culb.
chicitae Culb. [Cetraria] = Cetrelia chicitae (Culb.) Culb. & C.Culb.
chlorophylla (Willd.) Vainio [Cetraria] = Tuckermannopsis chlorophylla
(Willd.) Hale
chrysantha Tuck. [Cetraria] = Asahinea chrysantha (Tuck.) Culb. & C.Culb.
ciliaris Ach. [Cetraria] = Tuckermannopsis ciliaris (Ach.) Gyelnik
citrina Taylor [Cetraria] =? Nephromopsis pallescens (Schaerer) Park
clarkti Tuck. [Cetraria] {nomen nudum} = Nephromopsis endocrocea Asah.
f. clarktt Asah.
collata (Nyl.) Miill.Arg. [Cetraria] = Cetrelia collata (Nyl.) Culb. & C.Culb.
commixta (Nyl.) Th.Fr. [Cetraria] = Cetraria commixta (Nyl.) Th.Fr.
complicata Laurer [Cetraria] = Nephromopsis laureri (Krempelh.) Kurok.
coralligera (W.Weber) Hale [Cetraria] = Tuckermannopsis coralligera
(W.Weber) W.Weber
corallophora Mill.Arg. [Cetraria] =Punctelia rudecta (Ach.) Krog
corniculata Egeling [Cetraria] = Coelocaulon aculeatum (Schreber) Link
corrugis Fr. [Cetraria] = Parmotrema perforatum (Wulfen) Hale
crispa (Ach.) Nyl. [Cetraria] = Cetraria ericetorum Opiz
cucullata (Bellardi) Ach. [Cetraria| = Allocetraria cucullata (Bellardi) Randl.
& Saag
culbersonit Hale [Cetraria] = Cetraria culbersonii Hale
culbersoniorum ‘Trass [Asahinea] = Asahinea chrysantha (Tuck.) Culb. &
C.Culb.
daibuensis Rasanen [Cetraria] = Nephromopsis laxa (Zahlbr.) Sato
. davidiana Culb. & C.Culb. [Cetrelia] = Cetrelia davidiana Culb. & C.Culb.
. delavayana Culb. & C.Culb. [Cetrelia] = Cetrelia delavayana Culb. & C.Culb.
. delavayi (Hue) Sato [Cetraria] = Nephromopsis delavayi Hue
. delisei (Bory ex Schaerer) Nyl. [Cetraria] = Cetraria delisei (Bory ex
Schaerer) Nyl.
. denticulata Hue [Cetraria]
. dermatoidea Zahlbr. [Cetraria]
diffusa (Weber) Lynge [Cetraria] = Imshaugia placorodia (Ach.) S.F.Meyer
. ectocarpisma (Hue) Gyelnik [Nephromopsis] = Nephromopsis ectocarpisma
(Hue) Gyelnik
elenkinii Krog [Cetraria] = Cetraria nigricascens (Nyl. in Norrl.) Elenkin
398
S87
34.
DD:
36.
oe
*38.
39.
40.
41.
42.
*43.
44,
*40:
46.
*A7.
48.
49,
50.
endocrocea (Asah.) Sato [Cetraria] = Nephromopsis endocrocea Asah.
endoxantha Hue [Cetraria] = pro parte Nephromopsis endocrocea Asah., pro
parte Nephromopsis ornata (Miill. Arg.) Hue
endoxanthoides Awasthi [Cetraria] = Nephromposis endoxanthoides (Awasthi)
Randl. & Saag
epiphorella Nyl. in Crombie [Cetraria] = Coelopogon epiphorellum (Ny)l. in
Crombie) Brusse & Karnef.
ericetorum Opiz [Cetraria] = Cetraria ericetorum Opiz
eriophylla (Knight) Zahlbr. [Cetraria] = Erioderma knighti Shirley
erosa Culb. & C.Culb. [Platismatia] = Platismatia erosa Culb. & C.Culb.
everniella (Nyl.) Krempelh. [Cetraria] = Allocetraria stracheyi (Bab.) Kurok.
& Lai
fahlunensis sensu Schaerer [Cetraria] = Cetraria hepatizon (Ach.) Vainio
fahlunensis sensu Vainio [Cetraria] = Cetraria commixta (Nyl.) Th.Fr.
fallax (Weber) Ach. [Cetraria] = Platismatia glauca (L.) Culb. & C.Culb..
fastigiata (Delise ex Nyl. in Norrl.) Karnef. [Cetraria] = Cetraria fastigiata
(Delise ex Nyl. in Norrl.) Karnef.
fendleri (Nyl.) Tuck. [Cetraria] = Tuckermannopsis fendleri (Ny1.) Hale
formosana Zahlbr. [Cetraria] = Platismatia formosana (Zahlbr.) Culb. &
C.Culb.
frostii Tuck. [Parmelia] = Cetraria culbersonii Hale
furcellata Fr. [Cetraria] = Bryoria furcellata (Fr.) Brodo & D.Hawksw.
gilva Asah. [Cetraria| = Tuckermannopsis gilva (Asah.) Lai
glauca (L.) Ach. [Cetraria] = Platismatia glauca (L.) Culb. & C.Culb.
globulans (Nyl. ex Hue) Zahlbr. [Cetraria] = Nephromopsis globulans (Nyl.
ex Hue) Lai
gracilenta (Krempelh.) Vainio [Cetraria] = Coelopogon epiphorellum (Nyl. in
Crombie) Brusse & K4rnef.
halei Culb. & C.Culb. [Cetraria] = Tuckermannopsis americana (Sprengel)
Hale
hepatizon (Ach.) Kurok. [Tuckermannopsis] = Cetraria hepatizon (Ach.)
Vainio.
herrei Imsh. [Cetraria] = Platismatia herrei (Imsh.) Culb. & C.Culb.
hiascens (Fr.) Th.Fr. [Cetraria] = Cetraria delisei (Bory ex Schaerer) Nyl.
hypotrachyna Mill.Arg. [Cetraria] = Cetraria hypotrachyna Mill.Arg.
iberica Crespo & Barreno [Cetraria] = Tuckermannopsis merrillii (Du Rietz)
Hale
idahoensis Essl. [Cetraria] = Esslingeriana idahoensis (Ess}.) Hale & Lai
inermis (Nyl.) Krog [Cetraria] = Cetraria inermis (Nyl.) Krog
inflata J.D.Hook. & Taylor [Cetraria] = Ramalina inflata (J.D.Hook. &
Taylor) J.D.Hook. & Taylor
interrupta Culb. & C.Culb. [Platismatia] = Platismatia interrupta Culb. &
C.Culb.
isidiata (Asah.) Culb. & C.Culb. [Cetrelia] = Cetrelia isidiata (Asah.) Culb.
& C.Culb.
isidiigera Kurok. & Lai [Allocetraria] = Allocetraria isidiigera Kurok. & Lai
S79
51. isidioidea (Rasaénen) Awasthi [Cetraria} = Nephromopsis _ tsidioidea
(Rasénen) Randl. & Saag
52. japonica Zahlbr. [Cetraria] = Cetrelia japonica (Zahlbr.) Culb. & C.Culb.
53. islandica (L.) Ach. [Cetraria] = Cetraria islandica (L.) Ach.
54. juniperina (L.) Ach. [Cetraria] = Vulpicida juniperinus (L.) J.-E.Mattsson &
Lai
55. kamczatica Savicz [Cetraria] = Cetraria kamczatica Savicz
56. komarovii Elenkin [Cetraria] = Nephromopsis komarovii (Elenkin) Wei
kurodakensis Asah. [Cetraria] = Asahinea scholanderi (Llano) Culb. &
C.Culb.
*57. kurokawae Shibuichi & Yoshida [Cetraria] = Nephromopsis kurokawae
(Shibuichi & Yoshida) Kurok.
lacera J.D.Hook. & Taylor [Cetraria] = Sticta lacera (J.D.Hook. & Taylor)
Mill. Arg.
58. lacunosa Ach. [Cetraria] = Platismatia lacunosa (Ach.) Culb. & C.Culb.
lacunosa Elenkin [Cetraria] = Asahinea chrysantha (Tuck.) Culb. & C.Culb.
lacunosa Th.Fr. [Cetraria] = Platismatia norvegica (Lynge) Culb. & C.Culb.
laeteflava Zahlbr. [Cetraria] =? Cetreliopsis rhytidocarpa (Zahlbr.) Lai
59. laevigata Rassad. [Cetraria] = Cetraria laevigata Rassad.
lanata (Necker) Schaerer [Cetraria] = Pseudephebe pubescens (L.) M.Choisy
60. laureri Krempelh. [Cetraria] = Nephromopsis laurert (Krempelh.) Kurok.
*61. laxa (Zahlbr.) Sato [Cetraria] = Nephromopsis laxa (Zahlbr.) Sato
*62. leucostigma Lev. [Cetraria| = Cetraria leucostigma Lev.
libertina Stuckenb. [Cetraria] = Cetraria ericetorum Opiz
madreporiformis (Ach.) Miill.Arg. [Cetraria] = Dactylina madreporiformis
(Ach.) Tuck.
magnussonii Llano [Cetraria] = Cetraria nigricascens (Nyl. in Norrl.) Elenkin
megaleia (Nyl.) Rasanen [Cetraria] = Cetrelia sanguinea (Schaerer) Culb. &
C.Culb.
*63. melaloma Krempelh. [Cetraria] = Cetraria melaloma Krempelh.
*64. merrillii Du Rietz [Cetraria] = Tuckermannopsis merrillii (Du Rietz) Hale
265. microphylla Elenkin [Cetraria]
66. microphyllica Culb. & C.Culb. [Cetraria] = Tuckermannopsis microphyllica
(Culb. & C.Culb.) Lai
67. monachorum Zahlbr. [Parmelia] = Cetrelia monachorum (Zahlbr.) Culb. &
C.Culb.
68. morrisonicola Lai [Nephromopsis] = Nephromopsis morrisonicola Lai
muricata (Ach.) Eckf.[Cetraria] = Coelocaulon muricatum (Ach.) Laundon
69. nepalensis Awasthi [Cetraria] = Cetraria nepalensis Awasthi
nephromoides (Nyl.) Vainio [Cetraria] = Nephromopsis ectocarpisma (Hue)
Gyelnik
70. nigricans Nyl. [Cetraria] = Cetraria nigricans Ny].
71. nigricascens (Nyl. in Norrl.) Elenkin [Cetraria] = Cetraria nigricascens (Ny).
in Norrl.) Elenkin
nipharga (Ach.) R6ohling = Allocetraria cucullata (Bellardi) Randl. & Saag
72. nipponensis (Asah.) Culb. [Cetraria] = Nephromopsis nipponensis (Asah.) Lai
400
TS
nivalis (L.) Ach. [Cetraria] = Allocetraria nivalis (L.) Randl. & Saag
normoerica (Gunn.) Lynge [Cetraria] = Cornicularia normoerica (Gunn.) Du
Rietz
. norvegica (Lynge) Du Rietz [Cetraria] = Platismatia norvegica (Lynge) Culb.
& C.Culb.
nova-zelandiae Zahlbr. [Cetraria]
. nuda (Hue) Culb. & C.Culb. [Cetrelia] = Cetrelia nuda (Hue) Culb. &
C.Culb.
. oakesiana Tuck. [Cetraria] = Tuckermannopsis oakesiana (Tuck.) Hale
ochrocarpa (Eggerth) Lettau ([Cetraria} {nomen invalidum} =
Tuckermannopsis oakesiana (Tuck.) Hale
. odontella (Ach.) Ach. [Cetraria] = Cetraria odontella (Ach.) Ach.
. olivetorum Nyl. [Parmelia] = Cetrelia olivetorum (Nyl.) Culb. & C.Culb.
. orbata (Nyl.) Fink [Cetraria] = Tuckermannopsis orbata (Nyl.) Lai
. orientalis Randl. & Saag [Cetrelia] = Cetrelia orientalis Randl. & Saag
. omata Miill.Arg. [Cetraria] = Nephromopsis ornata (Miill.Arg.) Hue
pachysperma (Hue) Zahlbr. [Cetraria] = Parmelaria thomsonii (Stirton)
Awasthi
. pallescens Schaerer [Cetraria] = Nephromopsis pallescens (Schaerer) Park
pallida Awasthi [Cetraria] = Cetraria melaloma Krempelh.
. pallidula Tuck. ex Riddle [Cetraria] = Tuckermannopsis pallidula (Tuck. ex
Riddle) Hale
perstraminea Zahlbr. [Cetraria] = Nephromopsis komarovii (Elenkin) Wei
. pinastri (Scop.) Gray [Cetraria] = Vulpicida pinastri (Scop.) J.-E.Mattsson &
Lai
placorodia (Ach.) Tuck. [Cetraria] = Imshaugia placorodia (Ach.) S.F.Meyer
platyna Ach. [Cetraria] = Cetraria islandica (L.) Ach.
platyphylla Tuck. [Cetraria] = Tuckermannopsis platyphylla (Tuck.) Hale
. platyphylloides (Asah.) Sato [Cetraria] = Tuckermannopsis platyphylloides
(Asah.) Lai
polyschiza (Nyl.) Jatta [Cetraria] = Cetraria hepatizon (Ach.) Vainio
. potaninit Oxner [Cetraria| = Allocetraria potaninii (Oxner) Randl. & Saag
. pseudocollata Randl. & Saag [Cetrelia] = Cetrelia pseudocollata Randl. &
Saag
. pseudocomplicata Asah. [Cetraria] = Nephromopsis pseudocomplicata (Asab.)
Lai
. pseudolivetorum Asah. [Parmelia| = Cetrelia pseudolivetorum (Asah.) Culb.
& C.Culb.
pubescens (L.) Desport. [Cetraria] = Pseudephebe pubescens (L.) M.Choisy
ramulosa (Hook.) Tuck. [Cetraria] = Dactylina ramulosa (Hook.) Tuck.
. rassadinae Makryi [Cetraria] = Cetraria rassadinae Makryi
. regenerans Culb. & C.Culb. [Platismatia] = Platismatia regenerans Culb. &
C.Culb.
reticulata Krempelh. [Cetraria] {nomen nudum} = Platismatia erosa Culb. &
C.Culb.
rhizophora (Vainio) Rassadina [Cetraria] = Cetraria nigricascens (Nyl. in
Norrl.) Elenkin
*04.
95.
96.
se
98.
AMEE
100.
101.
102.
103.
104.
CALEY
*106.
Elie
*108.
109.
401
rhytidocarpa Mont. & v.d.Bosch [Cetraria] = Cetreliopsis rhytidocarpa (Mont.
& v.d.Bosch) Lai
richardsonii Hook. in Richardson [Cetraria] = Masonhalea richardsonii
(Hook. in Richardson) Karnef.
rubescens (Th.Fr.) Vainio [Parmelia] = Cetrelia olivetorum (Nyl.) Culb. &
C.Culb.
rugosa (Asah.) Sato [Cetraria] = Nephromopsis rugosa Asah.
sanguinea Schaerer [Cetraria] = Cetrelia sanguinea (Schaerer) Culb. &
C.Culb.
saviczii Oxner & Rassad. [Cetraria] = Asahinea scholanderi (Llano) Culb. &
C.Culb.
scholanderi Liano [Cetraria] = Asahinea scholanderi (Llano) Culb. & C.Culb.
scutata (Wulf.) Poetsch [Cetraria] = Tuckermannopsis sepincola (Ehrh.) Hale
scutata auct. [Cetraria] = Tuckermannopsis chlorophylla (Willd.) Hale
sepincola (Ehrh.) Ach. [Cetraria] = Tuckermannopsis sepincola (Ehrh.) Hale
septentrionalis (Nyl.) Almq. [Cetraria] = Asahinea chrysantha (Tuck.) Culb.
& C.Culb.
sibirica Magnusson [Cetraria] = Cetraria nigricascens (Nyl. in Norr].) Elenkin
sikkimensis Rasénen [Cetraria] = Cetraria leucostigma Lev.
simmonsii Krog [Cetraria] = Cetraria andrejevii Oxner
sinensis Culb. & C.Culb. [Cetrelia] = Cetrelia sinensis Culb. & C.Culb.
sorediella (Lettau) Rico & Manrique [Cetraria] {nomen nudum} = Cetraria
commicta f. sorediella Lettau
sphaerosporella Miill.Arg. [Parmelia] = Ahtiana sphaerosporella (Miill.Arg.)
Goward
stenophylla (Tuck.) G.K.Merr. [Cetraria] = Platismatia stenophylla (Tuck.)
Culb. & C.Culb.
stracheyi Bab. [Evernia] = Allocetraria stracheyi (Bab.) Kurok. & Lai
stracheyi Bab. [Cetraria] = Nephromopsis stracheyi (Bab.) Miuill.Arg.
straminea Krempelh. [Cetraria]
straminea Vainio [Cetraria] {nom. illegit} =? Cetreliopsis rhytidocarpa (Mont.
& v.d.Bosch) Lai
stuppea Flotow ex Sandst. = Coelocaulon muricatum (Ach.) Laundon
stygia (L.) Schaerer [Cetraria] = Melanelia stygia (L.) Essl.
subalpina Imsh. [Cetraria] = Cetraria subalpina Imsh.
subfendleri Essl. [Cetraria] = Cetraria subfendleri Ess\.
subperlatum Nyl. [Platysma] = Cetrelia sanguinea (Schaerer) Culb. & C.Culb.
subscutata Lindsay [Cetraria] = Cetraria subscutata Lindsay
subthomsoni Awasthi [Parmelaria] = Parmelaria subthomsonii Awasthi
sulphurea Mont. [Cetraria] =? Nephromopsis pallescens (Schaerer) Park
tenuifolia (Retz.) R.H.Howe [Cetraria] = Cetraria ericetorum Opiz
tenuissima (L.) Vainio [Cetraria] = Coelocaulon aculeatum (Schreber) Link
terrestris (Schaerer) Fink [Cetraria] = Vulpicida tilesti (Ach.) J.-E.Mattsson &
Lai
teysmanni Mont. & v.d.Bosch [Cetraria] =? Nephromopsis pallescens
(Schaerer) Park
402
110. thomsonii (Stirton) Miill.Arg. [Cetraria] = Parmelaria thomsonii (Stirton)
Awasthi
thyreophora (Ach.) Rohl. [Cetraria] = Cetraria islandica (L.) Ach.
111. tilesii Ach. [Cetraria] = Vulpicida tilesii (Ach.) J.-E.Mattsson & Lai
*112. togashii Asah. [Cetraria] = Cetraria togashit Asah.
tristis (Weber) Fr. [Cetraria] = Cornicularia normoerica (Gunn.) Du Rietz
113. tubulosa (Schaerer) B.de Lesd. [Cetraria] = Vulpicida tubulosus (Schaerer)
J.-E.Mattsson & Lai
tuckermanii Herre [Cetraria] {nom. illegit.} = Platismatia herrei (Imsh.) Culb.
& C.Culb.
tuckermanii R.H.Howe [Cetraria] {nom. illegit.} = Tuckermannopsis mernillii
(Du Rietz) Hale
114. tuckermanii Oakes [Cetraria] = Platismatia tuckermanii (Oakes ) Culb. &
C.Culb.
ulophylla (Ach.) Rebent. [Cetraria] = Tuckermannopsis chlorophylla (Willd.)
Hale
*115. ulophylloides Asah. [Cetraria] = Tuckermannopsis ulophylloides (Asah.) Lai
urceolata (Eschw.) Mont. [Cetraria] = Parmotrema sp.
116. viridis Schwein. [Cetraria] = Vulpicida viridis (Schwein.) J.-E.Mattssom & Lai
wallichiana (Taylor) Lai [Ahtia - nom. illegit.] = Cetrariopsis wallichiana
(Taylor) Kurok.
117. wallichiana (Taylor) Miill.Arg. [Cetraria] = Cetrariopsis wallichiana (Taylor)
Kurok.
*118. weberi Essl. [Cetraria] = Cetraria weberi Essl.
2119. xizangensis Wei & Jiang [Cetraria]
120. yunnanensis Zahlbr. [Cetraria] = Nephromopsis yunnanensis (Zahlbr.) Rand.
& Saag
ACKNOWLEDGEMENTS
We wish to express our gratitude to Dr.Ingvar Karnefelt, University of Lund,
for general support and reviewing this paper. Professor Teuvo Ahti (University of
Helsinki), Dr.Ming-Jou Lai (Dept. of Landscape Architecture, Fu-jen University,
Taipei), Jan-Eric Mattsson and Arne Thell (University of Lund) are thanked for
fruitful discussions.
LITERATURE CITED
Awasthi, D.D. 1987. A new position for Platysma thomsonii Stirton. Journ. Hattori
Bot. Lab. 63: 367-372.
Culberson,W.L. & Culberson, C.F. 1965. Asahinea, a new genus in the
Parmeliaceae. Brittonia 17: 182-190.
403
Culberson, W.L. & Culberson, C.F. 1968. The lichen genera Cetrelia and Platismatia
(Parmeliaceae). Contr. U.S. Natl.Hern.: 449-558.
Egan, R.S. 1987. A fifth checklist of the lichen-forming, lichenocolous and allied
fungi of the continental United States and Canada. Bryologist 90: 77-173.
Goward, T. 1985. Ahtiana, a new lichen genus in the Parmeliaceae. Bryologist 88:
367-371.
Gyelnik, V. 1931. Additamenta ad cognitionem lichenum extraeuropaeorum. Ann.
Cryptog. Exot. 4: 166-174.
Gyelnik, V. 1933. Lichenes varii novi critique. Acta Fauna et Flora Univ. 1: 3-10.
Karnefelt, I. 1977. Masonhalea, a new lichen genus in the Parmeliaceae. Bot.
Notiser 130: 101-107.
Kurokawa, S. 1980. Cetrariopsis, a new genus in the Parmeliaceae, and its
distribution. Mem.Natn. Sci. Mus. 13: 139-142.
Kurokawa, S. & Lai, M.J. 1991. Allocetraria, a new lichen genus in the Parmeliacea.
Bull. Natn. Sci. Mus. 17: 59-65.
Lai, M.J. 1980. Studies on the cetrarioid lichens in Parmeliaceae of East Asia.
Quart. Journ. Taiwan Museum 33: 215-229.
Mattsson, J.-E. & Lai, M.-J. 1993. Vulpicida, a new genus in Parmeliaceae
(lichenized Ascomycetes). Mycotaxon 46: 425-428.
Miller Argoviensis, J. 1891. Lichenes Miyoshiani in Japonia a cl. Miyoshi lecti et
a cl. Professore Yatabe communicati. Nuov. Giorn. Bot. Ital. 23: 120-131.
Rassadina, K.A. 1950. Cetraria in the U.S.S.R. Plantae Cryptogamae, 5: 171-304 (in
Russian).
Rasanen, V. 1952. Studies on the species of the lichen genera Cornicularia, Cetraria
and Nephromopsis. Kuopion Luonnon Ystavain Yhdistyksen Julkaisuja B 2:
1-53.
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MYCOTAXON
Volume XLVI, pp. 405-410 April-June 1993
TWO NEW SPECIES OF CRISTINIA
(BASIDIOMYCOTINA, APHYLLOPHORALES)
AND A SURVEY OF THE GENUS
Kurt Hjortstam
Malaregatan 12
S-441 35 Alingsas, Sweden
and
Helga Grosse-Brauckmann
Weingartenstrasse 10
D-6104 Seeheim-Jugenheim, Germany
Two new species of Cristinia Parm. are described viz. C. brevicellularis Hjortst. and
C. rhenana Grosse-Brauckmann. A new combination Cristinia coprophila (Wakef.)
Hjortst. is proposed. The genus Dacryobasidium Jiilich is considered as a synonym of
Cristinia. A key to the species of Cristinia is offered.
INTRODUCTION
Cristinia Parm. with the generic type Hydnum helveticum Pets., is a genus well character-
ized by arather loose basidiome, occurrence of hyphal strands and above all with cyanophi-
lous granulation in the basidia and thick-walled, cyanophilous basidiospores. The genus
was originally introduced with two species, the second being C. sasae Parm. which,
however, was later synonymized with C. helvetica by Baici and Hjortstam (1984).
CRISTINIA Parm. Consp. syst. corticiacearum p. 47 (1968).
BASIDIOMES resupinate, effused,loosely attached to the substratum.
HYMENOPHORES hypochnoid, mucedinioid, smooth to grandinioid, more rarely odontioid
or raduloid, hyphal strands occur regularly in the periphery.
- HYPHAL SYSTEM monomitic; hyphae thin-walled or with slight wall thickening, mostly
short-celled at least in the subhymenial layer, with or without clamp-connexions or
sometimes with scattered clamps on the subicular hyphae.
CYSTIDIA absent.
BASIDIA fairly short (15—25 tum), subclavate to cylindrical, slightly constricted, with four
sterigmata and a distinct cyanophilous granulation in the protoplasm.
SPORES fairly uniform, almost globose to subglobose and often slightly angular, 4-7 um
diam., smooth, moderately thick-walled to distinctly thick-walled and cyanophilous, not
amyloid or dextrinoid.
KEY TO SPECIES OF CRISTINIA
i. Clanip-Connexions absenigiegs ni tas send alte wy te eae PRN nu 2
1. Clamp-connexions present throughout, scattered or absent on the subicular hyphae 3
2. Hymenophore distinctly grandinioid, basal hypae 4—6 [um wide. Europe.
Be So PIG) nen RR cy ep Oe Vat a ieee PAR ER nach LN fit OY C. artheniensis
406
2. Hymenophore smooth to slightly grandinioid, basal hyphae 8-10 tm wide. Brazil.
ete Bl a gre eee rere ee mre te! oT ae eee bet NY Sete na C. brevicellularis
3. Hymenophore raduloid to irpicoid. An uncommon species. Holarctic .. C. gallica
32 Hymenophore smoothito erandinigid 92 eye, ee eee eee eee eee 4
4. Hymenophore grandinioid, all hyphae with clamp-connexions, spores 4.5—-5 um
broad. A rather common species. Cosmopolitan (7?) ........... C. helvetica
4. Hymenophore for the most part smooth, basal hyphae either without or with scat-
tered clamp=connexions, estat eta Pes AV cee sR eG oo aera ae ates 5
5. Hymenophore normally with a violet tint in KOH, basal hyphae with scattered clamp-
connexions, spores 5—7 um diam. Known from Germany only ....... C. rhenana
5. Hymenophore without a violet reaction in KOH, basal hyphae without clamp-con-
nexions, spores 4-5 um diam. England, additional records notknown . C. coprophila
SPECIES
1. CRISTINIA ARTHENIENSIS Baici & Hjortst. in Sydowia, Annls Mycol. Ser.II, 37:11
(1984).
2. CRISTINIA BREVICELLULARIS Hjortst. nov. sp.
Basidioma resupinatum, plus minus laxe adnatum, farinosum vel hypochnoideum. Hyme-
nium fere leve vel leviter grandinioideum, pallide isabellinum. Margo simillimus. Systema
hyphale monomiticum; totis hyphis efibulatis. Hyphae basales rectae, laxe intertextae,
tenuitunicatae, plus minus hyalinae, 8-10 um latae; hyphae subhymeniales brevicellulares,
5-8 tm latae. Hypharum fila presentes; hyphis tenuitunicatis vel crassiusculis, pallide
ochraceis, 5—8 tim latis. Cystidia nulla. Basidia subcylindracea, leviter constricta, 15-18 x
6-7 m, 4 sterigmatibus. Sporae subglobosae, leves, moderate crassitunicatae, circiter 5 x 4
m, neque amyloideae, neque dextrinoideae, distincte cyanophilae.
Holotypus: Brazil, Sao Paulo, Santos, Cananeia, Ilha do Cardoso, on deciduous wood, 2-5
Feb. 1987, Hjortstam 16753 (K).
Isotypus: (GB).
BASIDIOME resupinate, loosely adnate, farinaceous to hypochnoid.
HYMENOPHORE smooth to slightly grandinioid, pale isabelline. Hyphal strands present in
the periphery, with individual hyphae 5—8 um wide, pale ochraceous. Margin indeterminate.
HYPHAL SYSTEM monomitic; basal hyphae straight, branched at more or less right angles,
thin-walled, hyaline, 8-10 um wide; subhymenial hyphae short-celled, almost isodiametric
and with septa at 5-15 um intervals, 5-8 im wide; all hyphae without clamp-connexions.
CYSTIDIA absent.
BASIDIA subcylindric, with a distinct cyanophilous granulation, slightly constricted in the
middle part, 15-18 x 6-7 m, with four sterigmata, lacking a basal clamp-connexion.
SPORES subglobose, moderately thick-walled (KOH), smooth, about 5 x 4 m, inamyloid,
indextrinoid, strongly cyanophilous.
This is a typical species of Cristinia and is superficially similar to C. artheniensis Baici &
Hjortst. which also lacks clamp-connexions. The latter species, however, has a grandinioid
hymenophore and narrower hyphae.
407
3. CRISTINIA COPROPHILA (Wakef.) Hjortst. nov. comb.
Basionym: Corticium coprophilum Wakef. in Trans. Br. mycol. Soc. 6:480 (1916).
Holotype: England. On dung, beds near Temp. House, Kew, 5 July, 1912, Wakefield (K)!
This species is characterized by a nearly smooth and pale yellow hymenophore with hyphal
strands in the periphery. The basal hyphae are without clamp-connexions, but the sub-
hymenial ones regularly bearing clamps. The basidia are about 20-25 x 5.5—6.5 m, more
or less subclavate and with a distinct cyanophilous granulation. All basidia are provided
with a basal clamp-connexion and the spores are 4—5 im diam., thick-walled and distinctly
cyanophilous.
The species was reported by Jiilich (1972 and 1984) to be distributed in England
(type-locality), Germany, The Netherlands, Sweden and also in Canada and USA. Except
for the type no other specimens have been examined. The specimen from Sweden (Smaland,
Almesakra) represents Byssocorticium lutescens Erikss. & Ryv. and is the holotype of this
species and preserved in TRTC!
For illustration of Byssocorticium coprophilum and comments of B. lutescens see
Eriksson and Ryvarden (1973).
Jiilich (1981) described the genus Dacryobasidium, with Corticium coprophilum Wakef.
as type besides Byssocorticium lutescens Erikss. & Ryv. The latter lacks the basidial
cyanophilous granulation, and the morphology of the basidia as well as that of the spores
is more reminiscent of Hypochniciellum Hjortst. & Ryv. Also Amphinema Karst. and
Piloderma Jiilich should be considered, as the basidia are basally narrowed and more or
less pedunculate. The species is, however, at present maintained in Byssocorticium. Con-
sequently, the genus Dacryobasidium Jiilich is here considered as a synonym of Cristinia
Parm.
4. CRISTINIA GALLICA (Pilat) Jiilich in Persoonia 8:298 (1975)
Radulum gallicum Pilat in Mykologia 2:54 (1925); Cristinia mucida Erikss.& Ryv. nom.nov. Corticiaceae
North Eur. 3:311 (1975).
Radulum mucidum (Pers.) sensu Bourd. & Galz.in Bull. trimest. Soc. mycol. Fr. 30:247—248 (1914); non
Hydnum mucidum Pers.:Fr. Syst. Mycol. 1:418 (1821) = Dentipellis fragilis (Pers.:Fr.) Donk; non Hydnum
mucidum Pers. Mycol. Europ. 2:179 (1825) = Trechispora farinacea (Pers.:Fr.) Liberta.
For illustration and description (sub Cristinia mucida) see Eriksson and Ryvarden (1975).
See further Jiilich (op.cit.) for a nomenclatural discussion about Cristinia mucida Erikss.&
Ryv.
5. CRISTINIA HELVETICA (Pers.) Parm. in Consp. syst. corticiacearum p. 48 (1968)
Hydnum helveticum Pers. Mycol. Eur. 2:184 (1825).
For description and illustration of the species see Eriksson and Ryvarden (op.cit.).
6. CRISTINIA RHENANA Grosse-Brauckmann nov. sp. fig. 1
Basidioma resupinatum, laxe adherens, cremeum. Hymenium leve; margine fibrilloso cum
rhizomorphis subtilibus. Systema hyphale monomiticum, hyphae subhymeniales omnes
fibulatae, tenuiter tunicatae, 3.5—-5 tm latitudine. Hyphae basales fibulatae vel efibulatae,
latitudine 4-7(—9) m. Cystidia nulla. Basidia subclavata vel subcylindrata, (20—)25—35(—S0)
x (6—)6.5—7.5 m, 4 sterigmatibus. Protoplasma basidiorum cum guttulis oleosis cyanophilis.
408
Fig. 1. Cristinia rhenana. a) section through part of the basidiome b) basidia c) young
basidia with cyanophilous granulation d) spores. — From the holotype.
409
Sporae globosae vel obovatae, nonnumquam leviter angulatae, leves, cyanophilae, neque
amyloideae, neque dextrinoideae, incrassate tunicatae, diametro (4—)5—6.5(—7) m.
Holotypus: Germany, Hessen, Oberrheinebene, Nature Reserve “Kiihkopf’, on Populus
nigra, 8 Jan. 1988, H. Grosse-Brauckmann 3696 (GB).
Paratypi: Germany, Hessen, Oberrheinebene, Nature Reserve “Kiihkopf’, on Populus nigra,
31 Dec. 1988, H. Grosse-Brauckmann 4351; Rheinland-Pfalz, Oberrheinebene near Mann-
heim, ““Maudacher Bruch”, on Acer, 28 Dec. 1987, leg. U. Sauter. Duplicate H. Grosse-
Brauckmann 4371. All specimens in Grosse-Brauckmann’s priv. Herb.
BASIDIOME resupinate, effused, loosely attached to the substratum, soft, when fresh whitish
grey, in dried state cream-coloured, with thin hyphal strands in the subiculum and in the
periphery, margin indistinctly fibrillose.
HYMENOPHORE at first reticulate, then continuous and smooth, in KOH normally with a
violet tint.
HYPHAL SYSTEM monomitic; subhymenial hyphae indistinctly short-celled, thin-walled,
with clamp-connexions at all septa, 3.5-5 um wide; subicular hyphae somewhat thick-
walled and with scattered, mostly large clamp-connexions, 4—7(—9) um wide.
CYSTIDIA absent.
BASIDIA subclavate to subcylindrical, (20—)25—35(—50) x (6—)6.5—7.5 m, with oily contents
in the protoplasm and normally with four sterigmata, with a basal clamp-connexion.
Immature basidia with cyanophilous granulation.
SPORES globose to ovoid, now and then somewhat angular, with a small, but distinct
apiculus, smooth, thick-walled, inamyloid, indextrinoid, but distinctly cyanophilous,
mostly uniguttulate, (4-)5—6.5(—7) um diam.
The species is characterized by an almost smooth basidiome and scattered clamp-con-
nexions on the subicular hyphae. Further, the violet tint with KOH is distinctive.
SPECIES EXCLUDENDAE
Cristinia filia (Bres.) Liberta in Can. Journ. Bot. 51:1891 (1973); Corticium filium Bres. in
Annls mycol. Berlin, 6:43 (1908).
Lectotype: France, Bourdot no. 6984 (P), designated by Liberta (op. cit.).
This is not a species of Cristinia because of the acyanophilous basidial contents and the
- dextrinoid spores. Nevertheless, we have no suggestion as to its generic position.
Cristinia sonorae Nakas. & Gilb. in Mycologia 70:271—272 (1978).
The holotype, (USA, Arizona, KKN 237), is mixed and comprises two different species,
but neither belonging in Cristinia. One part is a species of Hyphodontia (subsectio
Ellipsosporae Parm.). This portion of the holotype fits the diagnosis of C. sonorae by
presence of cystidia and the size of spores, but neither hyphae nor spores are cyanophilous.
The second part seems to be a species of Radulodon Ryv. and is somewhat reminiscent of
R. americanus Ryv.
ACKNOWLEDGEMENTS
We wish to express our appreciation to Dr. Roy Watling, Royal Botanic Garden, Edinburgh
who revised the English language and to Prof. Leif Ryvarden, University of Oslo for
reviewing the manuscript. Additional thanks to Dr. Karl-Henrik Larsson, University of
Géteborg for editing and formatting the final manuscript.
410
REFERENCES
Baici, A. and Hjortstam, K. 1984. A new species of Cristinia (Aphyllophorales, Corticiaceae) from
North Italy. Sydowia Ann. Mycol. 37:1 1-14.
Eriksson, J. and Ryvarden, L. 1973. The Corticiaceae of North Europe, Vol. 2. Aleurodiscus —
Confertobasidium. Fungiflora. Oslo.
__ 1975. The Corticiaceae of North Europe, Vol. 3. Coronicium — Hyphoderma. Fungiflora. Osio.
Jiilich, W. 1972. Monographie der Athelieae (Corticiaceae, Basidiomycetes). Willdenowia, Beih.
7:1-283.
— 1981. Higher Taxa of Basidiomycetes. Bibliotheca Mycologica 85, 485 pp.
__ 1984. Die Nichtblitterpilze, Gallertpilze und Bauchpilze (Aphyllophorales, Heterobasidiomycetes,
Gastromycetes). Kleine Kryptogamenflora Band II b/1. Stuttgart New York.
MY COTAXON
Volume XLVI, pp. 411-413 April-June 1993
COMMENTS ON RECENT WORK ON
OPHIOSTOMA AND ITS SYNNEMATOUS ANAMORPHS
H. P. Upadhyay
Departamento de Micologia, Universidade Federal de Pernambuco
50739 Recife, Pernambuco, Brasil
The following is my reaction to two recent publications on the anamorphs of
Ophiostoma: Wingfield, M. J., Kendrick, B. and van Wyck, P. S. "Analysis of
conidium ontogeny in anamorphs of Ophiostoma." Mycological Research 95: 1328-
1333, 1991, and Mouton, M., Wingfield, M. J. and van Wyck, P. S. "Conidium
development in the synnematous anamorphs of Ophiostoma." Mycotaxon 46: 371-379,
1993. As I understand, the objective of these contributions were to study the type
species of Pesotum Crane & Sckoknecht and of Phialographium Upadhyay & Kendrick
_ developmentally in order to compare their conidiogenesis with that which occurs in the
type species of Graphium Corda. I am satisfied that the analysis that the authors
carried out was appropriate to answer the specific question that the authors posed.
However, I feel that this question itself is not the most important to raise regarding
the taxonomic value of percurrently proliferating conidiogenous cells in three genera,
since it is now apparent that the conidiogenous cell growth can no longer be viewed
as compartmentalized (Upadhyay, 1981; Minter et a/., 1983; Minter, 1987).
Although the authors entitled their paper "Analysis of conidium ontogeny in
anamorphs of Ophiostoma: Pesotum and Phialographium are synonyms of Graphium,"
they delineated, with some justification only two species of Pesotum, i.e. P. ulmi
(Schw.) Crane & Schoknecht and P. piceae Crane & Schoknecht as assigned by Crane
and Schoknecht (1973) vs. Graphium, but failed to examine the type species of
Phialographium. The rest of their discussions and decisions with respect to other
anamorphs of Ophiostoma were apparently based on scrutiny of the literature without
examination of types or authentic specimens. In my, A monograph of Ceratocystis and
Ceratocystiopsis (Upadhyay, 1981, p. 61), I cited a number of cultures and specimens
of the Phialographium state of Ceratocystis sagmatospora Wright & Cain (=P.
sagmatosporae Upadhyay & Kendrick) which they claimed could not be obtained for
EM studies. In my studies, these cultures of the type species of Phialographium
produced anamorphic and teleomorphic states, including conidiogenous cells with
distinct collarettes. I wish also to mention that the Hyalopesotum Upadhyay &
Kendrick, Pesotum, Pachnodium Upadhyay & Kendrick and Verticicladiella Hughes
anamorphs of Ceratocystis and Ceratocystiopsis, in my observations under phase
contrast photomicroscope, were commonly accompanied by a _ Sporothrix-like
morphogenesis in pure culture, either consistently or under certain conditions. Such
morphogenesis was never observed in the species of the form-genera Graphilbwm
412
Upadhyay & Kendrick, Leptographium Lagerberg & Melin, Phialocephala Kendrick,
or Phialographium that I examined. Ceratocystis ips (Rumb.) C. Moreau produced a
Graphilbum anamorph accompanied with Bainieria-like morphogenesis in most of the
cultures examined. On the other hand, synnematous and penicillately branched
mononematous conidiophores often lose fertility in artificial cultures to such an extent
that they may produce conidia only on simple conidiophores similar to those of
Sporothrix Hektoen & Perkins (Pesotwm-like and Verticicladiella-like) and Bainieria
Arnaud. These monomematous synanamorphs and their conidiation are important
issues that also need to be considered relative to the value of anamorph-generic
delimitation posed by the authors. Strangely enough, conidiogenous cells of Pesotum-
like and Verticicladiella-like anamorphs only can be discerned as_ proliferating
percurrently by the use of an electron microscope, which may not be feasible for every
worker for quick identification of species of these fungi.
Also, I would like to note that the first of these two articles contains two
erroneous citations: On page 1331 paragraph 2, the authors state, "Upadhyay and
Kendrick (1975) suggested that the conidiogenesis in the synnematous anamorph of
Ophiostoma sparsum (Davidson) de Hoog & Scheffer was phialidic, and established
the segregate anamorph-genus Graphilbum Upadhyay & Kendrick as the hyaline
analogue of Phialographium'"; whereas, Upadhyay and Kendrick (1975) clearly
characterized and delineated the genus Graphilbum as having percurrent, annellidic
conidiogenous cells (Mycologia 67: 800) and noted that this genus is a hyaline analogue
of Graphium (Mycologia 67: 799). Also, on page 1332, on the penultimate line, the
authors miscited Ceratocystis davidsonii Olchowecki & Reid, Can. J. Bot. 52: 1698
(1974) as "Ophiostoma davidsonii Olchowecki & Reid."
One reason that I feel that these issues need to be raised is that the use of
conidial ontogeny in delimitation of anamorphic genera is under intense scrtuiny
(Minter ef al. 1982, 1983, 1983, Minter, 1987). In light of present controversy
surrounding the use of conidial ontogeny in taxonomy of the ophiostomataceous fungi
and their anamorphic genera, the developmental study of types or authentic specimens,
preferably by comparing a combination of features of all closely related anamorphs and
teleomorphs, is necessary in order to understand and delineate the holomorph more
broadly than narrower definitions favored by Wingfield et al. For more detail, see
Upadhyay (1981, 1992)
Dr. Richard A. Humber, USDA-ARS, Ithaca, NY, critically reviewed that
manuscript, for which I am grateful. Dr. Leonor C. Maia of this Department kindly
read the manuscript.
LITERATURE CITED
Crane, J. L. and J. D. Schoknecht. 1973. Conidiogenesis in Ceratocystis ulmi,
Ceratocystis piceae, and Graphium penicilloides. Am. J. Bot. 60: 346-354.
Minter, D. W., P. M. Kirk and B. C. Sutton. 1982. Holoblastic phialides. Trans. Br.
Mycol. Soc. 79: 75-93.
Minter, D. W., P. M. Kirk and B. C. Sutton. 1983. Thallic phialides. Trans. Br.
Mycol. Soc. 80: 36-66.
413
Minter, D. W., B. C. Sutton and B. L. Brady. 1983. What are phialides anyway?
Trans. Br. Mycol. Soc. 81: 109-120.
Minter, D. W. 1987. The significance of conidiogenesis in pleoanamorphy. Pages
241-262 in: Pleomorphic fungi. J. Sugiyama, ed. Kodansha, Tokyo and
Elsevier, Amsterdam.
Upadhyay, H. P. and W. B. Kendrick. 1975. Prodromus for a revision of Ceratocystis
(Microascales, Ascomycetes) and its conidial states. Mycologia 67: 798-805.
Upadhyay, H. P. 1981. A monograph of Ceratocystis and Ceratocystiopsis. Univ.
Georgia Press: Athens, GA.
Upadhyay, H. P. 1993. Classification of the Ophiostomatalean Fungi. In: Ceratocystis
and Ophiostoma: Biology, Taxonomy, Ecology and Pathology. M. J.
Wingfield, K. A. Seifert and J. Weber, eds. American Phytopathological
Society Press. (in press).
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MYCOTAXON
Volume XLVI, pp. 415-424 April-June 1993
TREMATOSPHAERIA PACHYCARPA
AND HERBAMPULLA CRASSIROSTRIS
GEN. ET SPEC. NOV. (ASCOMYCETES)
Ch. SCHEUER & A. NOGRASEK
Institut fiir Botanik, Karl-Franzens-Universitat Graz
Holteigasse 6, A-8010 Graz, Austria
Botanamphora NOGRASEK & SCHEUER is a nomenclatural synonym
of Trematosphaeria FUCKEL. The unitunicate pyrenomycete described
by NOGRASEK & SCHEUER as Botanamphora pachycarpa (SACC. & El.
MARCHAL) NOGRASEK & SCHEUER is not identical with the type
material of Leptosphaeria pachycarpa SACC. & El. MARCHAL which
has bitunicate asci and belongs to the genus Jrematosphaeria. Both
fungi are redescribed and illustrated. Herbampulla crassirostris gen. et
spec. nov. is described to accommodate the unitunicate taxon.
Key words: Botanamphora pachycarpa, Leptosphaeria hazslinskyana,
Melanommataceae, Dothideales s.1., Melogrammataceae, Diaporthales s.].
The genus Botanamphora NOGRASEK & SCHEUER (in NOGRASEK 1990: 50)
is based on the type of Leptosphaeria pachycarpa SACC. & El. MARCHAL
which has been illustrated by BERLESE (1894: 78 and Tab. LXVI/3).
NOGRASEK & SCHEUER (l.c.) reported collections from the Eastern Alps
containing a very peculiar pyrenomycete with unitunicate asci which they
referred to Leptosphaeria pachycarpa. The type material of this taxon was
not available from PAD at that time, consequently the authors had to rely
on BERLESE 's drawings of ascus and ascospores for the new combination.
Trematosphaeria pachycarpa (SACC. & El. MARCHAL) SHOEMAKER &
BABCOCK (1989: 1595) is another combination based only on BERLESE's figure.
Furthermore, SHOEMAKER & BABCOCK synonymized Leptosphaeria
hazslinskyana BERLESE (1894: 78 and Tab. LXVII/1) with T. pachycarpa. So
far, the type material of L. hazslinskyana could not be located in any
herbarium.
416
The material of the SACCARDO collection in PAD has recently become
available on loan again. The type specimen of Leptosphaeria pachycarpa has
clearly bitunicate asci and the combination proposed by SHOEMAKER &
BABCOCK (1989) is most probably correct. Consequently, Botanamphora
becomes a taxonomic synonym of Trematosphaeria, and the unitunicate species
which has been misidentified as Leptosphaeria pachycarpa by NOGRASEK &
SCHEUER (l.c.) needs a new name.
Trematosphaeria pachycarpa (SACC. & El. MARCHAL) SHOEMAKER &
BABCOCK, Canad. J. Bot. 67: 1595 (1989).
Basionym: Leptosphaeria pachycarpa SACCARDO & El. MARCHAL, Rev.
Mycologique, Toulouse, 7: 145 (1885). |
Synonyms: Botanamphora pachycarpa (SACC. & El. MARCHAL) NOGRASEK &
SCHEUER in NOGRASEK, Biblioth. Mycol. 133: 51 (1990), sensu
SACC. & El. MARCHAL, non NOGRASEK & SCHEUER.
2? Leptosphaeria hazslinskyana BERLESE, Icones Fungorum I, p. 78
and Tab. LXVII/1 (1894).
Other descriptions (sub Leptosphaeria p.): SACCARDO, Syli. Fung. IX: 792
(1891); BERLESE, Icones Fung. I: p. 78 and Tab. LXVI/3 (1894).
Ascomata (pseudoperithecia) clustered or single, immersed to almost fully
erumpent, subepidermal, black, globose to laterally compressed, ca. 300-450
um in diam., with or without a rather broad, indistinct ostiolar papilla.
Peridium of variable thickness in the lower, immersed part of the ascoma,
often including + unaltered pieces of the substrate's sclerenchyma, consisting
of several layers of angular, sometimes slightly flattened cells of extremly
variable size and shape (cf. Fig. 1). In the erumpent part of the ascoma the
peridium is thicker, especially around the ostiole, with rather irregular,
blackish-brown cell wall thickenings in the outer layers. Interascal filaments
rather stiff, anastomosing (‘trabeculate’) up to 2.5 um thick in younger
ascomata with few mature asci, ca. 1 um thick between the asci of mature
ascomata, apparently embedded in a gel matrix, richly branching and loosely
interwoven in the upper part. Asci bitunicate (fissitunicate), eight-spored,
cylindric-clavate, ca. 140 x 16-18 um, seated on a basal layer of compact,
small-celled, hyaline plectenchyma inserted between the basal part of the
peridium and the centrum. The individual cells of the plectenchyma are
flattened in longitudinal section and apparently sometimes slightly interwoven.
Croziers not seen. Ascospores fusiform, ca. 40-52 x 7.5-10 um, S-septate
(2:3:1:3:2), brown with pale end cells; primary septum median.
417
Material examined: Leptosphaeria pachycarpa ‘SACC.’, in culmis graminum
majorum in Belgio (E1.
MARCHAL: Cryptogames de Belgique, Reliquiae
; Holotypus, PAD).
Westendorpi no. 86
The accommodation of Leptosphaeria pachycarpa in the genus Tremato-
sphaeria is accepted here because of the pallid end cells of the ascos
and the rather stiff, anastomosing (‘trabeculate’)
pores
and apically branched
interascal filaments which are apparently embedded in a gel matrix.
ypus,
Fig. 1. Trematosphaeria pachycarpa: Longitudinal section of ascoma (Holot
lactophenol permanent slide). Bar = 50 um.
418
Fig. 2 (@)
Fig. 2. Trematosphaeria pachycarpa: (A) Ascus. (B) Ascospores. (Holotypus,
lactophenol permanent slide). Bar = 20 um.
419
Herbampulla SCHEUER & NOGRASEK gen. nov.
Earlier description: sub Botanamphora NOGRASEK & SCHEUER in
NOGRASEK, Biblioth. Mycol. 133: 50 (1990).
Fam. ?Melogrammataceae, ord. Diaporthales
Typus generis: Herbampulla crassirostris SCHEUER & NOGRASEK, vide infra.
Etymology: lat. herba = grass (referring to the substrate)
lat. ampulla = bottle (referring to the shape of the perithecia);
literal translation of the Greek name Botanamphora.
Perithecia solitaria, in graminibus emortuis crescentia, substrato immersa,
ampulliformia. Peridium in parte externa (stromatica ?) paucis_ stratis
cellularum opacarum parietibus fuscis, in parte interna in sectione longitudinali
+ incoloratum, pluribus stratis cellularum applanatarum crassitunicatarum
guttulatarum compositum. Canalis ostioli periphysibus hyalinis, gelatinosis,
saepe in parte proximali furcatis vestitus. Peridium rostri intus textura oblita.
Paraphyses numerosissimae, filiformes, prae maturitatem ascorum +
degenerantes. Asci unitunicati, basibus deliquescentibus; annulus apicalis iodo
haud reagens, in sectione optica dua corpuscula refractivissima simulans.
Ascosporae + fusiformes, transverse septatae, maturae luteobrunneae, cellulis
terminalibus pallidioribus, guttulatae.
Perithecia single, immersed in the substratum, blackish brown, bottle-shaped.
Peridium with a conspicuous (stromatic ?) outer layer of small, dark brown
cells and a thicker inner layer of + colourless, guttulate cells which become
more thick-walled towards the outside. Ostiolar channel lined with numerous
hyaline periphyses, which are embedded in a gelatinous matrix and often
branched at the base, diverging inward from a textura oblita in the wall of
the neck. Paraphyses very numerous, filiform, + degenerated in mature
perithecia. Asci unitunicate, with a distinct, apical ring that appears as two
refringent bodies in optical section and shows no reaction to iodine, but is
stained deeply by Congo red. The bases of the asci deliquesce before
maturity and the asci are lying loosely in the perithecial cavity. Ascospores
+ fusiform, transversely septate, with pallid end cells, guttulate; primary
septum median.
Herbampulla crassirostris SCHEUER & NOGRASEK spec. nov.
Earlier description: sub Botanamphora pachycarpa (SACC. & El. MARCHAL)
NOGRASEK & SCHEUER in NOGRASEK, Biblioth. Mycol. 133: 51 (1990)
sensu NOGRASEK & SCHEUER, non SACC. & El. MARCHAL.
420
Etymology: lat. crassus = thick
lat. rostrum = beak (referring to the ostiole).
Perithecia substrato immersa, solum papilla ostioli ad 200 um longa
erumpentia, solitaria, rarius paulum gregaria, globosa vel ellipsoidea, fusca,
500-650 x 240-300 um. Papilla ostioli ad 190 um alta. Peridium 20-28 um
crassum, in parte externa (stromatica ?) paucis stratis cellularum opacarum
parietibus fuscis, in parte interna in sectione longitudinali + incoloratum,
pluribus stratis cellularum applanatarum guttulatarum compositum; extus
versus cellulae parietibus crassioribus, luminibus minoribus. Canalis ostioli
periphysibus hyalinis, gelatinosis, saepe in parte proximali furcatis, guttulatis
vestitus; peridium rostri intus textura oblita. Paraphyses filiformes, 2-5 um
crassae, in peritheciis maturis + degeneratae. Asci octospori, clavati-fusiformes,
80-129 x 10-12(-15) um, numerosisimi, annulo apicali conspicuo, ca. 2.8 x 2.8 um.
Ascosporae fusiformes, 30-44(-48) x 4-6(-7,5) um, extremis paulum
rotundatis, strictae vel leniter curvatae, 5-septatae, luteobrunneae, cellulis
terminalibus pallidioribus, guttulatae.
HOLOTYPUS hab. in foliis emortuis vix putrescentibus Caricis firmae [Carex
firma MYGIND): Austria, Tyrolia, in montibus ‘Lechtaler Alpen’ dictis, prope
oppidum 'Landeck’, in declivitatibus meridionalibus in vicinitate refugii 'Augs-
burger Hiitte’ supra pagum ‘Grins’, ca. 2200-2400 m, 09. VII.1982, leg. H.
Mayrhofer (GZU).
PARATYPE MATERIAL: On dead but not very decayed leaves of Carex firma
in the Caricetum firmae over calcareous substrates: AUSTRIA, Styria: Ausseer
Land, Totes Gebirge, ca. 6 km N of the lake Grundlsee, Redender Stein, ca.
1900 m, 08.VIII.1982, Ch. SCHEUER. - Ennstaler Alpen, Gesdéuse mountains,
southern ascent to the Admonter Reichenstein, near the fountain above the
Modlinger Hiitte, ca. 1700 m, 10.VIII.1985, Ch. SCHEUER. - Hochschwab moun-
tains, Trenchtling N of Trofaiach, little peak SE of the Hochturm, ca. 2000 m,
21.1V.1985, J. HAFELLNER. - Hochschwab mountains, peak of the GroBer Beil-
stein, NE of Aflenz, ca. 2000 m, 05.VIII.1984, J. HAFELLNER. - Hochschwab
mountains, peak of the Stangenwand, NE of Aflenz, ca. 2130 m, 04.VIII.1984, J.
HAFELLNER. - Eisenerzer Alpen, Reiting W of Trofaiach, NE slope of the
Grieskogel, ca. 2050 m, 09.VII.1984, J. HAFELLNER & A. NOGRASEK. - Eisen-
erzer Alpen, Polster, ca. 1900 m, 47°32'N/14°57'40"E, 15.VI.1984, W. OBERMAYER
& U. PRANTL (det. A. NOGRASEK). - On Sesleria varia (JAQU.) WETTST.:
SWITZERLAND, Graubiinden: Landwasser valley near Davos, Ziigenschlucht,
1300-1350 m, 28.VIII.1980, Ch. SCHEUER (microscope slide only).
Perithecia immersed in the substratum, with a prominent, thick, erumpent
beak up to 200 um in length, single, rarely in small groups, globose or
ellipsoid in outline, blackish brown, 500-650 x 240-300 um. Peridium 20-28 um
thick, consisting of a conspicuous (stromatic ?) outer layer of small, dark
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422
brown cells covered with dark brown hyphae, and a thicker, well
distinguishable, + colourless inner layer of slightly larger, more flattened,
guttulate cells. On the inside the cell walls are thin, towards the outside
they gradually become thicker, and the lumen smaller. Ostiolar channel lined
with hyaline, partly guttulate periphyses which are often forked at the base
and embedded in a gelatinous matrix, diverging inward from a textura oblita
in the wall of the neck. Paraphyses filiform, 2-5 um thick, + degenerated
in mature perithecia. Asci eight-spored, fusiform, 80-129 x 10-12(-15) um, very
numerous, apical ring ca. 2.8 x 2.8 um, staining deeply in a solution of Congo
red. Croziers not seen. Ascospores fusiform, 30-44(-48) x 4-6(-7.5) um, with
slightly rounded ends, 5-septate, yellowish brown with paler end cells,
guttulate (one larger oil drop in each cell).
At first, the monotypic genus Stioclettia DENNIS. (1975: 363, with St.
luzulina DENNIS) seemed to be quite appropriate for this peculiar fungus.
But a comparison of our taxon with the type material in K proved that there
are some important differences: (1) The ascus bases do not seem to dissolve
before maturity, which is also suggested by the narrowly cylindrical shape
of the asci with an uniseriate arrangement of ascospores. (2) The apical
apparatus as figured by DENNIS (1975) and visible in the permanent microscope
slides in K has a very different shape which reminds of a large Phomatospora.
(3) The ascospores are apparently colourless, not pigmented with paler end
cells. (4) The stromatic structures around the perithecia are probably more
prominent in Stioclettia, but the type material is rather poor and the
permanent microscope slides of longitudinal sections of the peridium in K
do not allow any other statement. It would not have been appropriate to
assign our new fungus to the wood-inhabiting, stromatic genus Melogramma FR.
either, although there is some similarity in spore structure, and probably
also in the apical apparatus of the asci and the development of the hymenium.
Therefore Herbampulla is tentatively placed in the Melogrammataceae.
Herbampulla crassirostris and Cainia graminis (NIESSL) v. ARX & MULLER
apparently prefer similar habitats and hosts. Both fungi occur on Carex firma
and Sesleria varia in the Caricetum firmae, but also in the Seslerio-Semper-
viretum on Sesleria varia, Cainia graminis also on Helictotrichon parlatorei
(WOODS) PILGER. Compared to other broad-leaved Gramineae and Cyperaceae
of the Alps, these plants have rather xeromorphic leaves and prefer relatively
well-drained localities - which also seems to be true for the small pyreno-
mycetes fruiting on their dead leaves in the early stages of decay.
Fig. 4. Herbampulla crassirostris: (A) Diagrammatic habit sketch of perithecia
in the substrate. (B) Asci and paraphyses. (C) Ascospores (Holotypus, lacto-
phenol permanent slides). (D) Ascospores (08.VIII.1982, Ch. SCHEUER; herba-
rium material in water). B-D: Bar = 20 um.
424
Acknowledgments
Thanks are due to Dr. Dietmar KORES for drawing our attention to
Herbampulla crassirostris, to Dr. Sergio CHIESA (Padova) for arranging the
loan of the type material of Leptosphaeria pachycarpa, to Prof. Dr. Lennart
HOLM and PD. Dr. Orlando PETRINI for acting as pre-submission reviewers.
References
BERLESE, A.N. (1894). Icones Fungorum omnium hucusque cognitorum.
Vol. I. - Avellino.
DENNIS, R.W.G. (1975). New or interesting British microfungi. III. - Kew
Bulletin 30: 345-365.
NOGRASEK, A. (1990). Ascomyceten auf GefaBpflanzen der Polsterseggen-
rasen in den Ostalpen. - Bibliotheca Mycologica 133: 1-271 + 5 pl. - Berlin,
Stuttgart: J. Cramer.
SACCARDO, P.A. (1891). Sylloge Fungorum omnium hucusque cognitorum.
IX. Supplementum universale. - Padova.
SHOEMAKER, R.A. & C.E. BABCOCK (1989). Phaeosphaeria. - Canadian
Journal of Botany 67: 1500-1599.
MY COTAXON
Volume XLVII, pp. 425-431 April-June 1993
NEW OR INTERESTING LICHENICOLOUS FUNGI. 3.*
KARSTENIOMYCES LLIMONAE SP. NOV. AND SCLEROCOCCUM
SERUSIAUXTI SP. NOV. (DEUTEROMYCOTINA)
Montserrat BOQUERAS
Dept. Biologia Vegetal (Botanica), Fac. Biologia Univ. Barcelona
Diagonal 645, E-08071 Barcelona, Spain
Paul DIEDERICH
Musée national d'histoire naturelle
Marché-aux-Poissons, L-2345 Luxembourg, G. D. Luxembourg
Abstract: Karsteniomyces llimonae Boqueras & Diederich sp. nov.
(Coelomycetes) is distinguished from related species by setose
pycnidia, much longer conidiophores and a different host (Parmelina
quercina); it is known only from the type locality in Spain
(Catalonia). Sclerococcum serusiauxii Boqueras & Diederich sp.
nov. (Hyphomycetes) is distinguished by very small sporodochia,
well-delimited thickened and darker regions of the conidial wall, and
the host (Parmelina pastillifera and P. tiliacea); it is known from
Spain (Navarra and Catalonia).
During an excursion in south Catalonia (Spain) in 1991, the authors collected an
appearently undescribed species of Karsteniomyces growing on Parmelina quercina.
A new species of Sclerococcum, growing also on species of Parmelina, has been
discovered by the senior author in Catalonia, and by Dr E. Sérusiaux and Dr J. Etayo
in Navarra (Spain). Both species are described below.
* 2. In: Bull. Soc. Nat. luxemb. 93: 155-162 (1992).
426
Karsteniomyces llimonae Boqueras & Diederich sp. nov.
Figs. (1-2).
Fungus lichenicola. Conidiomata pycnidia, singularia, dispersa, subglobosa
superficialia, ostiolata, straminea ad cinnamomea, 60-120 {zm diam., setis hyalinis,
non-septatis, 17-27 x 4-8 tum. Conidiophora elongata, simplicia ad sparse ramosa,
sparse septata, hyalina, 30-40 x 0.7-1 wm. Cellulae conidiogenae holoblasticae,
monoblasticae. Conidia sicca, elongato-ellipsoidea, hyalina, 1-septata, levia, 13-17 x
2.5-3.5 um.
Typus: Spain, Catalonia, Montsianés, La Sénia, Barranc del Retaule (Serra des
Ports), BF6914-BF6915, 1000-1100 m, on Corylus avellana, on Parmelina quercina,
10 11 1991, M. Boqueras, P. Diederich 9845 & A. Gémez-Bolea (BCC-Lich. 5224 -
holotypus; herb. Diederich - isotypus).
Conidiomata pycnidial, arising singly, scattered or loosely aggregated,
subglobose, superficial, with only the base entering the host tissues, ostiolate, pale
yellow orange to dark red orange, 60-120 {1m diam., with setae near the ostiole; setae
non-septate, hyaline, with a rounded apex, often constricted in the middle, 17-27 x 4-
8 um; pycnidial wall thin, composed of pseudosclerenchymatous cells, 15-25 tm
thick. Conidiophores arising from the inner pycnidial wall, cylindrical, elongate,
sometimes branched or septate, hyaline, 30-40 tum in length, 0.7-1 um thick.
Conidiogenous cells arising terminally and laterally from the conidiophores,
holoblastic, monoblastic, indistinguishable from the conidiophores except in the
absence of septa. Conidia dry, elongate-ellipsoid, with rounded ends or slightly
truncate at the base, hyaline, 1-septate, smooth-walled, with slightly unequal cells,
13-17 x 2.5-3.5 um.
Etymology: The species is named in honour of Dr Xavier Llimona (Barcelona,
Catalonia, Spain) for his enthusiasm and his contribution to the mycology (including
lichenology) of Catalonia.
Host: Parmelina quercina. The fungus acts as a parasite producing poorly
delimited decolorized patches on the host thallus.
Distribution: Known only from the type locality in south Catalonia (Spain).
Observations: The new fungus is easily distinguished from the two previously
known species of Karsteniomyces, K. peltigerae (P. Karsten) D. Hawksw. and K.
tuberculosus Alstrup & D. Hawksw., by the presence of hyaline setae near the
ostiole, by the much longer conidiophores, and by the different host. The two other
species of Karsteniomyces grow on Peltigera and both represent the anamorph of a
species of Scutula, S. miliaris (Wallr.) Trevis. and S. aggregata (Bagl. & Car.) Rehm
respectively.
427
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Because of the presence of setae near the ostiole and the much longer
conidiophores, the new species perhaps belongs to a different undescribed genus. As
long as no additional collections are known, we prefer, however, to include it in
Karsteniomyces.
The thin wall of most conidiomata of the original collection collapsed, giving the
conidiomata a cup-like appearance. Even with microtome sections, it was therefore
difficult to present a careful description and illustrations of the inner pycnidial wall
with the lower part of the conidiophores.
Sclerococcum serusiauxii Boqueras & Diederich sp. nov.
Figs. (3-4).
Fungus lichenicola. Conidiophora in sporodochiis densis convexis atrobrunneis,
rotundatis vel elongatis, (40-)60-180(-210) x (40-)60-100(-180) jum, semi-
macronemata, simplicia, atrobrunnea. Cellulae conidiogenae monoblasticae,
integratae, terminales, determinatae, ellipsoideae, atrobrunneae. Conidia variabilia, in
catenis irregularibus compactis, atrobrunnea, 9-20(-32) x 6-12 um, 2-14(-20) cellulis
subglobosis 3-7 [um diam., laevia, cum maculis obscurioribus.
Typus: Spain, Navarra, Col de Arette-Pierre-St-Martin, Larra, alt. 1550 m, in a
Pinus uncinata wood, on Parmelina pastillifera, 11 vii 1989, E. Sérusiaux 10604, J.
Etayo, P.W. James & F. Rose (LG - holotypus; herb. Diederich - isotypus).
Colonies forming convex dark brown to black, rounded or rarely elongate
sporodochia, (40-)60-180(-210) x (40-)60-100(-180) tum diam.; surface granular,
shiny; mycelium immersed in the host thallus. Conidiophores semi-macronematous,
not branched, agglomerated in very dense sporodochia, brown or reddish brown.
Conidiogenous cells monoblastic, integrated, terminal, determinate, ellipsoid to glo-
bose, reddish brown. Conidia variable in form, produced as irregular compact chains,
ellipsoid, clavate, sometimes lobed, dark brown, 9-20(-32) x 6-12 um, composed of
2-14-(20) subspherical or irregular cells of 3-7 {um diam., with a smooth surface, cell
wall unevenly thick, the thicker parts much darker than the rest of the conidium.
Etymology: The new species is named in honour of Dr Emmanuél Sérusiaux
(Li¢ge, Beigium) who collected the type specimen in the Pyrenees, and who allowed
us to describe it.
Hosts: Parmelina pastillifera and P. tiliacea (thallus, absent on the isidia). The
host thallus is not damaged by the presence of this fungus.
Distribution: Known from Navarra and south Catalonia (Spain).
Fig. 2. - Karsteniomyces llimonae (herb. Diederich - isotypus): Two pycnidia on
the thallus of Parmelina quercina; note the setae around the ostiole (x 80).
Fig. 3. - Sclerococcum serusiauxii (LG - holotypus): Thallus of Parmelina
pastillifera with numerous sporodochia (x 80).
430
Fig. 4. - Sclerococcum serusiauxii (LG - holotypus): Mature conidia, the two on
the left in optical section.
Observations: This species is easily distinguished from the two other known
Sclerococcum species with multicelled conidia by the very small sporodochia (they
are 170-350 tm in S. sphaerale (Ach.) Fr., and 225-500 um in S. epiphytorum
Diederich; see Diederich, 1990), by the granular surface of the sporodochia in which
conidia are often grouped in erect finger-shaped processes, by the larger number of
cells per conidium, and by the conidia showing uregular, well-delimited darker
regions due to an unevenly thickened cell wall, similar to that known in Milospium
graphideorum (Nyl.) D. Hawksw. (Hawksworth, 1975).
Additional specimens: Spain, Catalonia, Terra Alta, Ames, Barranc del Grevolar
(Serra des Ports), BF7125, 800-900 m, on Juniperus oxycedrus, on Parmelina
tiliacea, 20 x 1990, M. Boqueras, A. Farnés & A. Gémez-Bolea (BCC-Lich. 5225).
Spain, Navarra, Isaba, on P. tiliacea, s.d., Etayo s.n. (herb. Etayo).
431
Acknowledgements
We wish to thank Dr Emmanué] Sérusiaux (Liége) and Dr Javier Etayo
(Pamplona) for placing their collections of the new Sclerococcum at our disposal, Mr
Guy Marson (Luxembourg) for preparing microtome sections of Karsteniomyces
llimonae, as well as Prof. David L. Hawksworth and Dr E. Sérusiaux for reading and
commenting on our manuscript. The first author is obliged to the DGICYT, project nr
PB89-0518-C02-01, for the financial support.
References
Diederich P. (1990) - New or interesting lichenicolous fungi. 1. Species from
Luxembourg. Mycotaxon 37: 297-320.
Hawksworth D. L. (1975) - A revision of lichenicolous fungi accepted by Keissler in
Coniothecium. Trans. Br. mycol. Soc. 65: 219-228.
Hawksworth D. L. (1981) - The lichenicolous Coelomycetes. Bull. Br. Mus. nat. Hist.
(Bot.) 9: 1-98.
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Volume XLVI, pp. 433-437 April-June 1993
AMANITA REIDII - A NEW SPECIES FROM SOUTH AFRICA
ALBERT EICKER, J.V. VAN GREUNING AND DEREK A. REID
Department of Botany, University of Pretoria, Hillcrest, Pretoria 0002, South
Africa
SUMMARY
Amanita reidii is described and compared with the closely related A. phal-
loides (Fr.) Link
Since the publication of the recent monograph of South African species of Amanita
(Reid & Eicker, 1991) a further undescribed species has been found growing in
open stands of Eucalyptus. This new taxon is recognized by its grey-brown cap,
conspicuously streaked with darker radial markings and by its saccate volva. A
description of the new taxon follows:
Amanita reidii Eicker & Greuning, sp. nov.
(Figs 1-8)
Etym.: reidii - After Dr Derek A. Reid, for his significant contribution to South
African mycology
Pileus 3.0-7.2 cm diam., convexus vel applanatus, vel leviter campanulatus,
fusco-griseo-brunneus, centrum alibique versus pallide maculatescens; marginem
versus lineis conspicuis radialibus atrofuscis ornatus; superficies pilei glabra, sine
reliquiis veli; margo pilei non striatus. Lamellae adnexae, fere liberae, albae.
Stipes 3-7 cm altus, ad apicem 0.5-1.5 cm. diam., aequalis vel clavatus, basi usque
2.2 cm diam., albus vel sordide albus, annulo albo apicali membranaceo ornatus;
infra annulum superficie in squamas minutas fractiflexas disrupta. Volva libera,
alba, cylindrica ad stipitem arcte posita. Structura volvae hyphis angustis, usque 4
pm diam., efibulatis composita; nonnullis segmentis harum hypharum elongatis,
inflatis, usque 200 ym longis et 35 wm latis. Cheilocystidia 20-36 X 10-16 um,
clavata vel ovata parietibus tenuibus. Basidia 36-50 X 7-11 wm hyalina, clavata,
quadrispora parietibus tenuibus. Sporae subglobosae vel ovatae 7.5-8.0 X 6.0-
7.2, vel ovato-ellipticae 9.0-10.0 X 6.5-8.0 um, amyloidea, parietibus tenuibus;
sporae ex pulvere sporarum 8.0-9.5 X 6.2-9.0 wm metientes.
Holotypus sub Eucalyptus, Hideaway Game Farm, Melkrivier, prope
Pretoria, 29 April 1990; V. van Greuning & D.A. Reid. [in PRUM].
Pileus 3.0-7.2 cm convex or shallowly campanulate, becoming shallowly convex
or flattened, blotchily grey brown with an innately radially streaky surface. The
colour may vary to darker grey-brown or blackish slate-grey toward the centre and
become paler toward the margin, although the pileus often develops pallid dirty
white blotches at the disc or elsewhere. The streakiness is most conspicuous
toward the outer portion of the cap where the radial, dark grey-brown streaks con-
trast more sharply against the pale ground colour. __ Pileus lacking all traces of
velar tissue. Stipe 3-7 cm high, 0.5-1.5 cm wide at the equal or slightly enlarged
434
O
CoV OOD 3
Figs 1-7. Amanita reidii. 1. Spores from print mounted in Melzer's solution. 2.
Spores from gill mounted in Melzer's solution. 3. Basidia. 4. Elements from
universal veil. All from holotype. 5. Basidia. 6. Cheilocystidia. 7. Spores
from print mounted in NH, with congo red. All from Belfast collection. Bar =
10 pm. 2
apex, and up to 2.2 cm at the clavate, nonbulbous base, white to dirty white, the
surface with fibrillose scaly zig-zag markings below the annulus. Annulus apical,
pendulous, white, membranous. Volva white, saccate, and although free from the
stipe, sheathing it rather closely. | Lamellae adnexed to almost free, white to
creamy-white. Universal Velar tissue lacking from surface of pileus; structure of
volval limb hyphal, comprising narrow hyaline hyphae with a variable proportion
of very long inflated segments, but devoid of sphaerocysts; Occasional conducting
(gloeoplerous) hyphae present in some collections. The narrow hyphae, to 4 nm
wide, lack clamp connexions at the septa, and have thin but rather distinct walls.
The inflated segments, often fusoid, are usually of an intercalary position, up to
200 um long, and up to 35 wm wide. Partial veil hyphal. Cheilocystidia 20-36
X 10-16 wm thin-walled, hyaline clavate or ovate. In the Bergvliet material the
gill edge is formed of a thick layer of dissociating chains of thin-wailed, hyaline,
globose or ovate elements up to 30 um long and to 23 um wide. Basidia 36-50 X
7-11 pm, 4-spored, thin-walled. hyaline, clavate, originating from rounded or
435
wedge-shaped segments, but lacking a basal clamp-connexion. Spores from print:
8.0-9.5 X 6.2-9.0 wm (measured in Melzer's solution), varying subglobose to
ovate, more rarely ovate-elliptic, thin-walled, hyaline, amyloid. Spores taken
from the gill more variable in size and shape : measuring 7.0-8.0 X 6.0-7.2 ym
when subglobose or ovate, but 9.0-10.0 X 6.5-8.0 um when ovate-elliptic.
Distribution: under Eucalyptus claeziana, Bergvliet State Forest, Sabie,
G.C.A. van der Westhuizen (35), 7 Dec. 1984 [PREM 48618]; under Eucalyptus,
Hideaway Game Farm, Melkrivier, V. van Greuning & D.A. Reid, 29 April 1990
[PRUM 3179] [Holotype]; in open situation under Eucalyptus, Pineglades, Bel-
fast, 8 Feb. 1991. These gatherings are deposited in PRUM (Mycology Section of
the H.G.W.J. Schweickerdt herbarium, PRU, University of Pretoria) or PREM.
Edibility nothing is known regarding the edibility of this species, but be-
cause of a suggested relationship with A. phalloides it should be treated with very
great caution and avoided.
DISCUSSION
A. reidii is undoubtedly closely related to A. phalloides (Fr.) Link, differ-
ing in its prominently streaky grey-brown pileus, without trace of green tints. By
contrast the pileus of A. phalloides is yellow-green to olive-green, ornamented with
fine, innate, rather densely crowded and slightly darker radial streaks. In A. reidii
the streaks are fewer but darker and altogether more pronounced. The white stipe
of A. reidii also differs from that of A. phalloides, which is often greenish, in that
it is not conspicuously bulbous as in the latter species, and is rather closely
sheathed by the volva. Again A. reidii appears to be of slightly smaller stature.
Further the known collections of A. reidii have all comprised fruitbodies growing
in the vicinity of Eucalyptus trees, and it may be that this fungus will ultimately be
shown to require a more specialised habitat than does A. phalloides.
In South Africa there has been a trend toward naming the fungus described
above as A. phalloides f. umbrina Ferry - at least on field meetings. However the
name has not yet appeared in the South African literature.
Through the kindness of Dr. Jan Kuthan and Dr. Herink we have received
copies of Ferry's original description and comments on this taxon, from which it is
clear that he regarded his forma uwmbrina as representing the ultimate stage of
ageing of the familiar yellow, greenish or olive fungus. As such it does not merit
taxonomic recognition. Discussing the variation of A. phalloides, Ferry (1911)
wrote "A ces diverses variétés on pourrait ajouter une forme umbrina ....... Ce
n'est pas une variété; les diverses variétés jaune, verdatre ou olive peuvent par-
venir a ce stade, qui représenté une état plus avancé : cette teinte bistre survient
avant qu'il y ait la moindre trace d'altérative ou ni la moindre odeur."
Since we have observed specimens of the fungus under discussion in all
stages of development at the Melkrivier locality without finding any evidence of
green tints in the young fruitbodies, we feel confident that the South African
material represents an autonomous taxon distinct from both Amanita phalloides f.
phalloides and A. phalloides f. umbrina. We have, therefore, decided to publish
it at specific rank under the name A. reidii.
Reference to the publications of Bottomley and Talbot (1954) and van der
Westhuizen (1983) suggest that they were aware of this brown to grey brown
Amanita, but that they included it in their concept of A. phalloides (Fr.) Link.
Thus Bottomley & Talbot (1.c.) in their description of the latter referred to it as
having the cap "smooth or streaked with dark innate fibrils and to be variable in
colour from "greenish, yellowish olive, greyish brown, umber brown to smoky
olive, or even darker ......... ". Indeed their italics suggest that umber brown to
smoky olive was the more commonly met with colour form, and further suggests
436
that it was A. reidii rather than the true A. phalloides which was the more familiar
species to these authors - at least in the Transvaal. Van der Westhuizen (1983)
also noted that A. phalloides, in South Africa, occurred in various shades of
greenish yellow olive, smoky brownish-green, umber brown to smoky olive,
darker towards the centre." This description again appears to have embraced A.
reidii.
Having considered the taxonomic relationship between A. reidii and the
other species of Amanita known to occur in South Africa it is necessary to consider
this relationship in a wider context. It would seem that the South African species
is most closely related to A. alauda Corner & Bas (1962), which was described
from the Gardens Jungle, Botanic Gardens, Singapore, based on a single collection
comprising two half specimens. This species belongs in Amanita, subgenus
Lepidella, section Phalloideae, along with both A. phalloides and A. reidii.
Fig. 8. Amanita reidii. Two fruitbodies from Melkrivier collection x 1.
A. alauda is very similar to A. reidii in having a brownish cap devoid of
green colouration, varying from sepia or pale purplish umber to mouse-coloured,
darker at the centre. More importantly the cap is "streaked by innate dark fibrils,
especially near margin". However the stature of A. alauda differs from that of A.
reidii in that whereas the latter tends to be rather short and squat with the length of
the stem about equal to the diameter of the cap, A. alauda is taller and more
elegant in appearance with the length of the narrow stem approaching twice the
diameter of the cap.
Further the cap of A. alauda has a distinct umbo seated in a central depres-
sion; in contrast the cap of A. reidii is either shallowly campanulate with a flat-
tened, sometimes almost truncate disc, or applanate;. umbonate sporophores have
not been observed in this species. Another difference is that the cap of A. alauda
is more distinctly brown in colour, and the radial streakiness is less complete and
less obviously reticulate. Further the cap surface of A. alauda was described as
437
viscid, a feature emphasized in the authors discussion of their new taxon. A vis-
cid cap cuticle has not been observed in A. reidii. Again the surface of the stem
of A. alaudia was said to be smooth; in A. reidii it disrupts into a zig-zag pattern
of fibrillose markings, but this may not be of great taxonomic significance. Spore
size and shape are very similar in both taxa as are the basidial features.
Cheilocystidia are also similar although these organs are very slightly broader in A.
alauda. However comparison of the account of the microstructure of the volval
limb of A. alauda with that of A. reidii suggests that there are differences in this
respect. Corner & Bas (1962) wrote "Limb of volva mainly consisting of 3-5(-25)
pm wide, interwoven sublongitudinal hyphae, with some scarce sphaerocysts up to
100 um wide in the inner part". There is no mention of the occurrence of very
long, rather abruptly inflated fusoid hyphal segments such as those found in the
volval tissue of A. reidii. Finally there is the difference in habitat to be con-
sidered for while A. reidii appears to grow in association with Eucalyptus, which is
suggestive of a mycorrhizal relationship, A. alauda was said to grow in jungle with
the implication that it grew amongst native vegetation.
Finally since A. reidii has, to date, been found only in areas strongly in-
fluenced by human activity, it seems that this fungus will prove to be an exotic
species in South Africa. If so one might look to Australasia as its possible
geographical origin.
ACKNOWLEDGEMENTS
The authors are grateful to the South African Foundation for Research
Development for financial support. We thank the Director General, Department of
Water Affairs and Forestry for permission to collect fungi on their property.
REFERENCES
Bottomley, A.M. & Talbot, P.H.B. (1954). Common Edible and Poisonous
Mushrooms in South Africa. Bulletin 324, Botany and Plant Pathology
Series No.17, Department of Agriculture, Union of South Africa.
Corer, E.J.H. & Bas, C. (1962). The genus Amanita in Singapore and Malaya.
Persoonia 2: 241-304.
Ferry, R. (1911). Etudes sur les Amanites. A. phalloides, A. verna, A.virosa,
Revue mycologique. Toulouse. Suppl. 1.
Reid, D.A. & Eicker, A. (1991). South African fungi : the genus Amanita.
Mycological Research 95, 80-95.
Van der Westhuizen, G.C.A. (1983). Mushrooms and Toadstools. Bulletin 396.
Plant Protection Research Institute, Department of Agriculture, South
Africa.
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MY COTAXON
Volume XLVII, pp. 439-460 April-June 1993
PHYTOPHTHORA HIBERNALIS AND P. SYRINGAE
H.H. Ho’ and 8.C. Jong’
‘Department of Biology,
State University of New York,
New Paltz, New York 12465
*Mycology & Botany Department,
American Type Culture Collection,
12301 Parklawn Drive, Rockville, Maryland 20852
ABSTRACT
A comparative morphological study of isolates
of Phytophthora hibernalis and PP. syringae
confirmed that these are two distinct species,
readily distinguished by the easily deciduous,
elongated ovoid to ellipsoidal sporangia with long
pedicels and the amphigynous and paragynous
antheridia in varying proportions in the former and
the nondecidous, broadly ovoid, obpyriform to
ellipsoidal sporangia and the predominantly
paragynous antheridia in the latter. Confusions
in literature concerning these two species can be
attributable to misidentification and/or acceptance
of Tucker's treatment of P. hibernalis synonymous
with P. syringae which had priority.
INTRODUCTION
In 1881, Berkeley first described the fungus
attacking the leaves of lilac under the binomial
Ovularia syringae. The pathogen was later renamed
as the type of a new genus: Phloeophthora syringae
Kleb. (Klebahn, 1906), then Nozemia syringae
(Kleb.) Pethybr. (Pethybridge, 1913) and finally
as Phytophthora syringae (Kleb.) Kleb. (Klebahn,
1909), characterized. by sympodially developed
sporangiophores, large NOvoOLd, obovoid to
440
ellipsoidal" sporangium with a "flat cap at the
énd" sand the ~ productiion “of oogonia: with
paragynous antheridia. Despite Sarejanni's
attempt (1936) to treat it as a subvariety of P.
cactorum, P. syringae has been accepted widely
aS a good species.
In 1925, Carne reported a new species of
Phytophthora, P. hibernalis on the fruit, leaves
and smaller branches of Citrus spp. in Australia.
It was unique in its "very deciduous, ellipsoidal
or lemon shaped conidia with a constant pedicel"
and the production of oogonia with amphigynous
and occasionally paragynous antheridia. The
fungus wasS considered so different from P.
syringae that Carne did not even attempt to
compare these two species when he discussed the
systematic position of P. hibernalis. However,
the morphological similarity between P. syringae
and P. hibernalis was noted by Petri (1927) and
later by Tucker (1931) who showed that these two
species were identical in the production of only
paragynous antheridia on all culture media, their
failure to grow at 25 C, their growth characters,
the production of oogonia and oospores of almost
the same dimensions, and similarly small
sporangia with flattened papilla, sparse on agar
medium but more abundant in water. Consequently,
he concluded that P. hibernalis should be treated
as synonymous with P. syringae which has
priority. Leonian (1934) also found that these
two species were very similar both
morphologically and physiologically but he
preferred to keep them as separate species, an
opinion shared by Waterhouse (1963) and other
researchers. However, Wager (1942), Frezzi
(1950)5 Noviello and Snyder (1963) and
Novotelnova (1974) subscribed to Tucker's thesis
that P. hibernalis and PP. syringae are
conspecific. Ther di Chotomy eh bys Op Lntons
concerning the taxonomic status of these two
species often led to confusion in literature,
especially when both have been known to attack
the same host, for example, citrus during cool
weathers causing identical symptoms (Klotz and
Calavan, 1969). In a comprehensive study of the
441
sexual reproduction of Phytophthora species,
Savage et al. (1968) pointed out the urgent need
for a thorough study of isolates of P. hibernalis
and P. syringae. However, most of the studies
in the past centered on the pathology and ecology
of P. syringae, especially on lilac (De Bruyn
1924a,b; 1928), apple, pear and related Rosaceae
fruit trees (Harris, 1979, 85 a,b,c; Harris and
Cole, 1982; Upstone, 1978; Upstone & Gunn, 1978;
Doster and Bostock, 1988b; Young and Milbrath,
1959), sweet chestnut and beech (Day, 1938) and
P. hibernalis on citrus (Favaloro and Sammarco,
LO Fst LOt Ze ali 7 3) ¥
The present study was undertaken to compare
the morphological characteristics of various
isolates of P. hibernalis and P. syringae under
uniform conditions in order to define the species
more precisely.
MATERIALS AND METHODS
Isolates and media: Specific information on
the isolates of P. hibernalis and P. syringae
used is given in Table 1. With the exception of
N37 which was provided through the courtesy of
Dr. Robert Young, Department of Plant Pathology,
West Virginia University, Morgantown, Va., all
isolates were obtained from the American Type
Culture Collection (ATCC), Rockville, Maryland.
Unless otherwise stated, clarified V-8 juice agar
medium (Ribeiro, 1978) Supplemented with
sitosterol (30mg/1) (CV8) was used for culture.
Morphology: Colony characteristics on CV8
were compared after incubating the cultures in
darknessaeats) 20, Car foreu7ardays’. The colony
diameters were measured at right angles through
the inoculum, and the width of primary hyphae
measured using a light microscope. Sporangia
formation was induced by incubating under light
at 20C small mycelial agar discs on cCV8 in
freshly collected stream water that had been
filtered through 0.45 um pore size membrane
discs. Sex organs in single cultures were
examined periodically by microsocopy through the
442
bottom of the petri dish. If the isolate failed
to produce sex organs in single culture, it was
then grown on Tryptophane Medium (Ribeiro, 1978)
or cCV8 supplemented with 10% vegetable oil
(Doster and Bostock, 1988a) and incubated at 7-
SCs
RESULTS
Colony morphology
All isolates were characterized by rather
slow growth rate on cCV8 (4-8mm/day for P.
hibernalis and 4-6mm/day for P. syringae),
appressed to moderately fluffy, often showing
distinct growth patterns, commonly rosette (with
broad, rounded petalloid sectors) Les.
hibernalis and chrysanthemum (with small, narrow,
rounded petaloid sectors) in P. syringae. The
hyphae were narrow, 3-5 um wide for P. hibernalis
and 3-6 wm wide for P. syringae often irregular
and wavy with frequent short branches. Small
spherical to oval, monilioid hyphal swellings
(under 25 um) were commonly produced by some
isolates of P. syringae in water or in old
cultures but not by isolates of P. hibernalis.
No chlamydospores were produced by any isolate
under any experimental condition.
Sporangia
All isolates produced sporangia readily in
water within 24-48 hrs, except ATCC 60352 P.
hibernalis and ATCC 34002 P. syringae. Isolates
ATCC 56352 P. hibernalis and ATCC 14837 P.
syringae also produced sporangia on agar medium
though not as abundantly as in water. The
sporangia of P. hibernalis were rather small (av.
29-53 x 14-22 um), characteristically elongated
ovoid to ellipsoidal, often with tapered base and
the broadest part near the semi-papillate apex
(L/B av. 1.8-2.4), borne terminally on unbranched
or long branches of narrow (1-2 um wide), lax
sympodially branched sporangiophores and were
easily deciduous, each with a long pedicel (23-
73 wm). In contrast, the sporangiophores of P.
443
syringae were either unbranched or distinctly
sympodially branched bearing non-deciduous
sporangia terminally on short branches often with
conspicuous basal plugs and _ subsporangial
swellings. The semi-papillate sporangia were in
general larger (av. 41-75 x 39-40 um) with mostly
rounded base and rather wide and flat apex
(apical thickening 1-3 um), ovoid to ellipsoidal
in water (L/B av. 1.4-2.1) but broadly ovoid to
obpyriform on agar plate (L/B 1.5). No internal
proliferation of sporangiophore was observed in
any isolate. Nearly all sporangia released
zoospores in water through single exit pore, but
most sporangia of ATCC 56352 P. hibernalis
germinated directly by one or more than one germ
tubes through the apex whereas ATCC 14837 P.
syringae occasionally formed secondary sporangia
in a chain as demonstrated by Lafferty and
Pethybridge (1922) Noviello and Snyder (1963) and
Edney and Chambers (1981).
sex Organs
Most isolates produced sex organs in single
cultures either readily in 2-3 wks or tardily
atter} 2-—5yamontnhs. Isolates ATCC 60352 P.
hibernalis and ATCC 34002 P. syringae failed to
produce them under any condition. Two isolates
of P. syringae, ATCC 46886 and 64876 produced sex
organs sparsely only in old cultures but more
rapidly and abundantly on V8C supplemented with
vegetable oil. The oogonia of both species were
colorless to slightly yellowish to tawnish with
the single oospore nearly filling the oogonial
space, and the oospore wall averaged 1-3 um
thick. In general, the oogonia and oospores of
P. hibernalis were slightly smaller than those
of P. syringae (av. 33-35 um verses 33-41 um diam
and 29-32 um versus 30-35 um diam).
Whereas the antheridia were distinctly
predominantly paragynous in all isolates of P.
syringae, they were predominantly amphigynous in
isolate ATCC 64708 but variable in isolates ATCC
32995 and 56353 of P. hibernalis. The antheridia
were similarly small for both species, averaging
444
10 x 10 wm for P. hibernalis and 11 x 10 um for
P. syringae. None of the isolates studied here
produced chlamydospores.
DISCUSSION
Our study of various isolates of Phytophthora
hibernalis and P. syringae confirmed Waterhouse's
conclusion (1963) that these are two distinct
species. There is little doubt that they have
much in common, for instance, their homothallism,
their low maximal temperature for growth, their
narrow, wavy, irregular hyphae, small antheridia,
plerotic oospores and their often rosette
cultural patterns. However, P. hibernalis can
be easily distinguished from P. syringae
primarily by the deciduous, elongated ellipsoidal
and usually smaller sporangia with a tapered base
bearing a long pedicel. Most isolates of P.
syringae produced well-defined sympodially
branched sporangiophores with broadly ovoid to
obpyriform sporangia borne on short stalks and
in no instance were the sporangia deciduous. The
formation of small moniliform hyphal swellings
is a common feature of some P. syringae isolates.
These differences were also noted by previous
workers (Table 4). The antheridia of P. syringae
have been described as paragynous or
predominantly paragynous. Our study showed that
whereas both types were present, the paragynous
antheridia were undoubtedly far more common than
amphigynous antheridia. On the other hand, there
was much confusion regarding the antheridial
configuration in P. hibernalis which has been
described as amphigynous, predominantly
amphigynous, paragynous or predominantly
paragynous (Table 4). Gerrettson-Cornell (1989)
stated that the antheridia of P. hibernalis have
been reported to be predominantly amphigynous in
fresh cultures but predominantly paragynous in
old cultures. We have not been able to confirm
this in our literature research. In fact, we
noticed that in isolates ATCC 32995 and 56353 of
P. hibernalis most antheridia appeared initially
as paragynous but later on, amphigynous
antheridia often prevailed. It is quite possible
445
that the ratio of amphigynous and paragynous
antheridia produced in culture may be variable
as found in P. porri (Ho, 1983) and P. megasperma
(Erwin, 1965), thus explaining the discrepancies
in reports regarding the antheridial types of P.
hibernalis.
The confusions in literature about the
morphological characteristics of P. hibernalis
and P. syringae can also be attributable, at
least in some cases, to misidentification. Thus,
Nadel-Schiffmann (1947) reported and described
"Pp, hibernalis" from citrus in Palestine and this
paper was listed as a major reference in the
species description of P. hibernalis (Waterhouse
and Waterston, 1964b). Based on the information
and the photographs presented by Nadel-Schiffmann
in her paper, we agree with kKouyeas) and
Chitzanidis (1968) that the fungus should have
been identified as P. syringae. Savage (1963)
studied an orange isolate of "P. hibernalis"
(N37) received from Klotz in California, and
found many oogonia with amphigynous antheridia
on 3-month lima-bean agar slants. Brennemann
(1967) re-investigated N37 along with two other
isolates of "P. hibernalis" (N116 received from
CMI and N252 received from CBS) and reported that
they produced paragynous antheridia on hemp-seed
and oatmeal agar plates. He suspected too, that
these isolates might be P. syringae, but did not
attempt to study them thoroughly to determine
their real identity. We have been able to obtain
isolate N37 and based on the ~- sporangial
characteristics, there is little doubt that it
should have been assigned to P. syringae instead
of P. hibernalis.
Accepting Tucker's treatment of P. hibernalis
to be synonymous with P. syringae which has
priority, Wager (1942), Frezzi (1950) and Verneau
(1953) reported "P. syringae" on citrus even
though the pathogen had all or most of the
features characteristic of P. hibernalis as
described by Carne (1925). Wager described the
antheridia of " P. syringae" as paragynous, yet
his diagram of two oogonia with attached
446
antheridia showed one to be paragynous and the
other, in all likelihood, amphigynous, resembling
the amphigynous antheridia of P. cinnamomi and
P. parasitica in the same figure. It is puzzling
why Tucker (1931) considered P. hibernalis
identical as P. syringae. He compared single
lilac isolate of P. syringae from Baarn with
single isolate of P. hibernalis "from Kew to
Baarn" and found that both produced paragynous
antheridia in all media, nearly identical oogonia
and oospores and similar sporangia with flattened
papillae. There was no mention of the caducity
of sporangia in the isolate of "P. hibernalis" -
- a feature seldom missed by other researchers
(Table 4). It is not inconceivable that the "P.
hibernalis" isolate that Tucker examined was
actually P. syringae.
Once these possible sources of confusions
have been identified, the differences between
these two species reported in literature became
apparent. Thus, the sporangia of P. hibernalis
have been reported as easily deciduous,
ellipsoidal, averaging 37 (10-75) x 18 (8-35) um;
L/B 2.1 (1.8-2.5), with long pedicels (2-80 um),
closely matching our measurements in present
SCUGY 39°" 408 (29-55 LP xe, POR Gl 4220 eeu Meranate4 ee ee
(1.8-2-4), pedicel 23-73 um long. In contrast,
P. syringae produced non-deciduous, broadly
ovoid, obpyriform to elongated, larger sporangia,
averaging 4%. (271475) "xs 307@ 1-42) um Bae ©
(12-233) Vins lLiteraturetand, 547(41=75)ex"32,(24—
40) umeetis/iB yi8 Cle a—oeiye ine Our, Study. Both
species prefer low temperature for growth, with
the minimum at 0-5 C and maximum 22-27 C. The
antheridia of P. syringae are predominantly
paragynous. Those authors who reported the
antheridia simply as paragynous probably had
missed the amphigynous antheridia which could be
rare in some isolates. On the other hand, the
antheridia of P. hibernalis can best be described
as having amphigynous and paragynous antheridia
in varying proportions in view of the conflicting
reports in literature and based on our
observations. There is little difference in the
size of sex organs between these two species
447
based on published data. The oogonia, oospores
and antheridia of P. hibernalis and P. syringae
averaged, respectively, 23 (27-56) vs. 31 (18-
50) wm diam, 28 (20-40) vs. 28 (17-38) um diam
and 11.5 (9-21) x 11.5 (10-15) vs. 10 (10-19.6)
x 8 (7-10) um. It is interesting to note that
in our study, the oogonia and oospores of P.
syringae appeared slightly larger than those of
P. hibernalis, averaging respectively 37 (33-41)
vs. 34.(33-35) um diam and 33 (30-35) vs. 31 (29-
32) um diam. Our measurements of the antheridia
of P. hibernalis and P. syringae showed them to
bes -simplarnlyvasmalilsalOmex LOM VS 6 al ax y1O0eum.
Although both species produce petalloid colonies
described commonly as "rosette" many isolates of
P. syringae produced "chrysanthemum" type growth
patterns, with small narrow rounded sectors.
Chlamydospores are absent in P. hibernalis or P.
syringae. The chain-like "chlamydospores"
reported by Nadel-Schiffman (1947) for "BP.
hibernalis" should be interpreted as moniliod
hyphal swellings for the lack of delimiting septa
(Blackwell, 1949).
ACKNOWLEDGEMENTS
This work was supported in part by NSF Grant
DIRS 9O= 15 ls ie COmoGcd:. The authors thank Elmer
Davis and Dr. H.S. Chang for reviewing the
manuscript.
LITERATURE CITED
Al-Hedarthy & Tsao 1979. Sporangium pedicel
length in Phytophthora species and the
consideration of its uniformity in
determining sporangium caducity. Trans. Br.
MV.COLMES OChee/ 2c) lia F,
Barrachina-Tuset, J.J. 1977. Contribution al
conocimiento del genero Phytophthora de Bary
en Espana. Ann. Inst. Nacional de Invest.
Agrarias Ser. Prot. Veg. 7: 11-106.
Bensaude, M. 1929. Note sur le Phytophthora,
parasite des Citrus au Portugal. Compt. Rend.
soc. Biol. 101: 982-984.
Berkeley, M.J. 1881. Lilac fungus. Gard. Chron.
448
Toei 665%
Blackwell, E. 1949. Terminology in Phytophthora.
Mycol. Pap. 30. Commonw. Mycol. Inst., Kew,
Surrey, England. 24pp.
Boccas, B. and E. Laville, 1978. Les maladies
a Phytophthora, des agrumes. Inst. de
Recherches sur les Fruits et Agrumes. 162
Pp.
Bolay, A. 1977. Phytophthora syringae - agemt
d'une grave pourriture des pommes-~ en
conservation. Rev. Vitic. Arbor. Hort. 9:
LoL=169.%
Bostock, R.M. and M.A. Doster. 1985. Association
of Phytophthora syringae with pruning wound
cankers of almond trees. Pl. Dis. 69: 568-
Sorel
Brasier, C.M. and R.G. Strouts. 1978. New or
unusual records of plant diseases and pest.
Pl. Path. 27: 99-100.
Brenneman, J.A. 1967. Patterns of sexuality in
ten species of Phytophthora. M.S. Thesis,
W.Va. Unin::, Morgantown, W: Va. 9°53) pp”
Buddenhagen, I.W. 1955. Various aspects
Phytophthora cactorum collar-rot of apple
trees in the Netherlands. Tijdschr. Plziekt.
Le ES WE 28 ed te
Carne, W.M. 1925. AMeDY OWN TOC OLMmCTerus iin
Australia (Phytophthora hibernalis n. sp.)
J. (Roy. Soctun Wie Australivaivi2 713 —49".
Chester, R.S. 1932. A comparative study of three
Phytophthora diseases of Lilac and of their
pathogens. J. Arnold Arboretum 13: 209-232.
Day; W.R.2 1938.) *Root-rot of) sweet) chestnut and
beech caused by species of Phytophthora. J.
SOC VFOn.MGre (Bri Csr a1 O05.
De Bruyn, H.L.G. 1924a. The Phytophthora disease
of lilac. | Phytopathology 14" 503=5i7"
De’, Bruyn,’ H.L.Gs 29246; De oorzaak van het
epidermisch optreden van de Phytophthora
ziekte van de seringen. Tijdsch. Plziekt.
S30 DLS 12 24
De Bruyn, H.L.G. 1928. Is ontbladering als
bestrijidinswijze tegen Phytophthora z ziekte
van de seringen gewenscht? Tijydsch':?..Pr*
ziekt. 34: 233-238.
Doepel, R.F. 1966. Phytophthora species on
449
citrus in Western Australia. Pl. Dis. Reptr.
50: 494-496.
Doster, M.A. and R.M. Bostock. 1988a. The effect
of temperature and type of medium on oospore
production by Phythphthora syringae.
Mycologia 80: 77-81.
Doster, M.A. and R.M. Bostock. 1988b. Chemical
protection of almond pruning wounds from
infection by Phytophthora syringae. Pl. Dis.
72: 492-494.
Edney, K.L. and D.A. Chambers. 1981. Post-
harvest treatments for the control of
Phytophthora syringae storage rot of apples.
ANN. wappLemBroley O07 si 23 7-241.
Erwin, D.C. 1965. Reclassification of the causal
agent of root rot of alfalfa from
Phytophthora crytogea to P. megasperma.
Phytopathology 55: 1139-1143.
Favaloro, M. and G. Sammarco. 1973. Ricerche sul
marciume del collette e radicale degli
agrumi. Specie di Phytophthora presente
negli agruneti della Sicilia Orientale.
Phytopath. Medit. 12: 105-107.
Fawcett, H.S. 1933. New locations for
Phytophthora citrophthora and P. hibernalis
on citrus. Phytopathology 23: 667-669.
Fawcett, H.S. 1936. Citrus diseases and their
control. McGraw-Hill, New York. 656 p.
Frezzi, M.J. 191507. Las especies de
'Phytophthora' en la Argentina. Rev. Invest.
AgGreiGwiB.wAl Festacy 4421354
Gerrettson-Cornell, L. 1989. A compendum and
classification of the species of the genus
Phytophthora de Bary by the canons of the
traditional taxonomy. For. Comm. New South
Wales Tech. Pap. 45, Sydney. 103 p.
Harris, D.C. 1979. Epidemiology of Phythophthora
syringae storage rot. East Malling Res. Sta.
Rept.) 1978 5:-193-94°:
Harris) auD. Coun l9S5a. Survival of Phytophthora
syringae oospores in and on apple
erchardsoil:.. Trans wBru.wMyYcoln. Soc, 8 os 153=—
nies 5 a
Harris, D.C. 1985b. Microdochium fusarioides sp.
nov. from oospores of Phytophthora syringae.
Trans se pre MYVGOl. SOC. 84 58—36 17
450
Harris, D.C. 1985c. The colonization of fallen
apple leaves by Phytophthora syringae in
relation to inoculum levels in orchard soil.
Ann. Appi. “BLOLE. 1072 179=188.
Harris, D.C. and D.M. Cole. 1982. Germination
of Phytophthora syringae oospores. Trans.
Bre MYCOL. foOC es o..mipie) OV et
Himmelbaur, W. 1911. Zur kenntnis' der
Phytophthoreen. Jahrb. Hambur. Wiss. Anst.
28: 39-61.
Ho, H.H. 1983. Phytophthora porri from stored
carrots in Alberta. Mycologia 75: 747-751.
Klebahn, H. 1905. Eine neue Pilzkrankheit der
syringen. Centbl. Bakt. 15: 335-336.
Klebahn, H. 1909. Die neue Zweig-und
Knospenkrankheit. Krankheiten des Flieders.
Gebruder Borntralger, Berlin. 18-75.
KlOUZ, teri. Maloy se Color Handbok of Citrus
DiseasesscUnavs) OL Caltts Divs of pAGr=1SCc.
E227°ps
Klotz, L.J. and E.C. Calavan. 1969. Gum diseases
Of CTlUcrus. nie Caliitornia. Unive i Ofmca lite.
AGE. EXpve sta. #Clr mo 96. es ender eV. «2. Jan.
Kouyeas, H. and A. Chitzanidis. 1968. Notes on
Greek species of Phytophthora. Ann. Inst.
Phytopath.}Benaki, N.S. 8; 175-192.
Krober, H. 1985. Experiences with Phytophthora
de Bary and Pythium Pringsheim. Bundesanst
fur Land - und Forstw. (Berlin-Dahlem) 255:
A sei iy Yl 852
Lafferty, H.A. and G.H. Pethybridge. 1922. On
a Phytophthora parasitic on apples which has
both amphigynous and paragynous antheridia;
and an allied species which show the same
phenomenon’... Proc. "Roy. Dublin’ soc. 17: .29—
43.
Leonian, LH. 1934. Identification of
Phytophthora species. W.Va. Agric. Exp. Sta.
Bull. 262. 36 pp.
McLennany +E. [7 1935" Notes on the organisms
causing’ brown yrot Of = "citrusmetrure, ein
Victoria: Austraila (Phytophthora
Citropnthorare(sm-.) 7k) om). = Leon strata.
hibernalis Carne). Proc. Roy. Soc. Victoria
(N.S.) 48: 96-103.
Moniz, da Maia, R. 1925. Um nuovo Oomycete
451
parasita de a'rvores do genero Citrus? Rev.
Agron (Lisbon) eser.a4% jen —)1y
Nadel-Schiffmann, M. 1947. Phytophthora
hibernalis. Palestine J. Bot. Rev. Ser. 6:
148-157
Newhook, F.J., G.M. Waterhouse and D.J. Stamps.
1978. Tabular key to the species of
Phytophthora de Bary. Mycol. Pap.) 143),
Commonw. Mycol. Inst., Kew, Surrey, England.
20ND"
NOV1C1.1O wy Commer OnGQueeWsC san GONVOC) mie lO6G. A
Phytophthora disease of fennel. Phytopath.
zeitsch. 46: 139-169.
Novotel 'Nova, N.S. 1974. The Genus Phytophthora.
Soviet Acad. Sci. Bot. Inst. Lening. U.S.~S-R.
206 p.
eyopukigk=s Say eslchke A fruit rot of apples and
pears due to a variety of Phytopthora
syringae. Long Ashton Agr. Hort. Res. Sta.
Annu. Rept. 1930: 147-150.
Peglion, V. and M. Sacchetti. 1927. Intorno alla
peronospora del Lilla (Phytophthora syringae,
Klebahn). Atti. R. Accad. Naz. Lincei, Rend.
Cl. Sci. Fis., Mat. e Nat. 5: 696-698.
Pethybridge, G.H. 1913. On the rotting of
potatoe tubers by a new species. of
Phytophthora having a method of sexual
reproduction hitherto undescribed. Sc. Proc.
Ree DLLD eS OCLs FL oe OA
Petri, L. 1927. Rasseyna del cai fitopatologia
pin notevoli osservat: nel, 1926. Bol. R.
S Cac baw. VeCO,.m/: lsd oepp.
Ribeiro, O.K. 1978. A Source Book of the Genus
Phytophthora. J. Cramer, Vaduz, Germany.
417 p.
Rosenbaum, J. 1917. Studies of the genus
Phyvtophtnora...J. Agnus (Res. (830 233-206.
Salih, H. 1974. A Phytophthora species new for
Turkey determined in citrus orchards in
Adanas Use lLULK. swrny.CODaCh 06:96) ja 1 5
Sarejanni, J.A. 1936. La pourriture du collet
des Solanees cultivees et la classification
du genre Phytophthora. Ann. Inst. Phytopath.
Benake 2:35-52.
Savage, 4b. UemmlooS . Sexuality in the genus
Phytophthora II. Intra- and interspecific
452
pairings. Ph.D. Thesis, W.Va. Univ.,
Morgantown, W.Va. 87 p.
Savage; E.J.,;,C.W. Clayton, J.H. Hunter,” J.-A.
Brenneman, C. Laviola and M.E. Gallegly.
1968. Homothallism, heterothallism and
interspecific hybridization in the genus
Phytophthora. Phytopathology 58: 1004-1021.
Schwinn, Pear ae hove ue Untersuchungen zur
systematik der gattung Phytophthora de Bary.
ArCHGMIKTODLOl. oS eco oo.
Sewell, G.W.F. and J.F. Wilson. 1964. Death of
maiden apple trees caused by Phytophthora
syringae Kleb. and a comparison of the
pathogen with P. cactorum (L. & C.) Schroet.
ANN. “ADDL. (DIGle oo... 200 ~2oUR
Stamps, D.J., G.M. Waterhouse, F.J. Newhook and
G.S. Hall. 1990. Revised tabular key to the
species of Phytophthora. Mycol. Pap. 162.
COA. B. 2INCern sy MVCOWs VANSG. 7) NeW Vo oULneyy
England. 28 °p
Tucker, Ce M wees. Taxonomy of the genus
Phytophthora de Bary. Mo. Agr. Exp. St. Res.
Bulla’: 153.0 208 ip.
Upstone, M.E. 1978. Phytophthora syringae fruit
rot) (ofmapples Vi PinyPath. 327 524-505
Upstone, M.E. and E. Gunn. 1978. Rainfall and
the occurrence of Phytophthora syringae fruit
rot of apples in Kent 1973-75. ‘Pl. Path. 27:
30=—3 5
Verneau, R. 1953. Le Phytophthora parasite degli
agruni nella Campania. Nota 1. Specie
osservate sui frutti. Annali Exp. Agaaria
Gems OG Oar.
Wager, V.A. 1942. Pythiaceous fungi on citrus.
Hilgardia 14: 535-548.
Waterhouse, G.M. 1963. Key to the species of
Phytophthora de Bary. Mycol... Paps. 92.
Commonw. Mycol. Inst., Kew, Surrey, England.
Ze Pp.
Waterhouse, G.M. and E.M. Blackwell. 1954. Key
to the species of Phytophthora recorded in
the British Isles. Mycol. Pap. 57. Commonw.
Myco..;/Inst.| Kew, Surrey,” England. 9. p,.
Waterhouse, G.M. and J.M. Waterston. 1964a.
Phytophthora syringae. Descriptions of
pathogenic fungi and bacteria 32. Commonw.
453
Mycol. Inst., Kew, Surrey, England. 2 p.
Waterhouse, G.M. and J.M. Waterson. 1964b.
Phytophthora hibernalis. Descriptions of
pathogenic fungi and bacteria 31. Commonw.
Mycol.) Inst. Kew, sourrey, England. 1i2 p.
VOUNG) eRe Ae aug eocmA. Mi LDA chm. S59 A stem
canker disease of fruit tree nursery stock
caused by Phytophthora syringae.
Phytopathology 49: 114-115. (Abstr. )
454
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MYCOTAXON
Volume XLVI, pp. 461-466 April-June 1993
A NEW CORTINARIUS FROM SOUTHEASTERN TEXAS
ORSON K. MILLER JR.
Department of Biology
Virginia Polytechnic Institute and State
University, Blacksburg, VA 24061
SUMMARY
INTRODUCTION
Southeastern Texas has large areas of mixed pine and
hardwood forest with loblolly pine (Pinus taeda L.)
intermixed with beech (Fagus grandifolia Ehrh.) and southern
oaks including swamp chestnut oak (Quercus prinus L.), sand
jack oak (Q. virginia v. geminata Sarg.), willow oak (Q.
phellos L.), and swamp red oak (Q. falcata var. pagodaefolia
Ell.). Watson (1979) describes several dominant plant
communities including areas of beech-magnolia-loblolly pine
forest, which is a transitional association between the
wetlands and the dry uplands. It occurs in the uplands and
hilly terrain interspersed with successional lakes or acid
bogs "where underground water seeps to the surface". In this
area Watson (1979) refers to the more advanced vegetation in
old lake beds as baygalls, which consist of sweet bay
(Magnolia virginiana L.) and gall berry holly (Ilex opaca
Ait.) along with black gum (Nyssa sylvatica var. biflora )
and red bay (Persea barbonia). These vegetational area are
often in close proximity to each other. Within this complex
plant community a rather abundant, undescribed, distinctive,
bright yellow Cortinarius first came to the attention of
David Lewis who made notes on the fresh characteristics,
habit and habitat. It is especially common along the boarder
of the baygall communities in the transition zone to the
higher elevation referred to as the beech-magnolia-loblolly
pine community. Since the baygall community has plants that
are typically endomycorrhizal, an ectomycorrhizal host must
be found elsewhere. David Lewis and I have observed
fruitings most often in association with beech, but loblolly
pine or the various oaks which are subdominant members of
the beech-magnolia-loblolly pine community might also be
associated.
Methods: Each collection was placed in a separate wrapper to
avoid mixing material. The possible mycorrhizal hosts were
recorded at the time of collection. Smell and taste were
recorded for specimens at the time of collection and again
in the laboratory. The color standards used included Ridgway
462
GLODZ5T, (the colors placed in quotes) and Kornerup and
Wanscher, (1978) (the colors indicated by (3A7) which in
page 3, row A and color block number 7). Microscopic
examination was carried out using 3% KOH and Melzer's
solution. All collections are deposited at VPI unless
other wise indicated.
TAXONOMY
Cortinarius lewisii sp. nov. O.K. Miller
Illustrations: Texas Mushrooms p.177.
Pileus: (1-) 2-3-5 cm latus convexus, plano-convexus
aut paene convexus umbone lato, squamulis aridis parvis
appressis; margo incurvatus luteus aut aurantius. Lamellae
adnexae aut adnatae confertae angustae seriebus duobus
Gispositae, in juventute albae, in maturitate aurantiacae,
in vestustate ferrugineae. Stipes 4-7.5 X 0.3-0.9 (-1.1) cm
latus aridus aequalis, fibrillis tenuibus juventute album,
luteum in vestutate, annulo luteo prope apicem in
maturitate. Odor moschatus aut raphano similis. Sapor
mitis. Sporae 6-7.5 X 5-6 um diam., subglobosae, verrucis
demissis brunneis. Depositum sporatum ferrugineum.
Pileipellis mixocutis est hypharum filamentosarum 3.4-8.4 um
diam., fibrillis praesentibus. Trama lamellaris parallela
hyalina parietibus crassiusculis.
Typum legit. D.P.Lewis-4370, 13 May 1990, Lance Rosier
Unit, Big Thicket National Preserve, Hardin Co., Texas.
Byleus:, (lO. e2-3 50cm -=broad pconvex;. Dlano-COnvex asco
nearly plane with a broad umbo in age, dry, with minutely
appressed, squamules, surface bright orange-yellow "Cadmium
Yellow", "Light Cadmium", margin "Apricot Yellow", buttons
chrome yellow (3A5), buttercup yellow, sunflower yellow
(4A8), to some cadmium orange (5A8). Margin incurved at
first, plane in age. Lamellae: adnexed to narrowly adnate,
close, narrow, with two tiers of lamellulae, nearly white in
buttonsto Pisabellat o(2.5Y.6.5/4) ;."Orange=-Citrine: ~sinvage
becoming rusty brown from the spores. Stipe: 4-7.5 cm long,
3-29 (-11.), mm; wide, dry, equal; coverediwithjrine fiprits,
"Light Cadmium" to cream color with white mycelium over the
base Partial veil: dense, cortinous, white in button but
soon yellow as pileus matures, forming a bright yellow
annulus near the apex but often absent at maturity. lesh:
firm, "Deep Colonial Buff", to "Isabella Color". ODOR: musky
to radish-musky. TASTE: mild.
Pileipellis: 80-180 yum thick, a mixocutis of filamentous
hyphae 3.4-8.4 um diam, loosely interwoven at the surface,
more compact below, with scattered intracellular
incrustations, occasional branching, scattered clamp
connections, yellow to orange in Melzer's solution, hyaline
in 3% KOH. Pileus trama: of broad hyphae 4-16 um diam,
becoming closely packed resembling a textura angularis in
463
Figs. 1-3. 1, 2 and 4 sp'd basidia and a basidiole. 2.
basidiospores. 3. filamentous clamped hyphae of the
pileipellis.
Scale bar = 10 umm.
464
the mid to lower trama, often with common, thick-walls (0.3-
0.6 wm) and scattered clamps, hyaline in 3% KOH, yellow in
Melzer's solution. Lamallar trama: parallel, thin-walled,
filamentous, hyaline, hyphae 2.5-7.6 um diam slightly thick-
walled near the pileus trama, often with scattered clamp
connections, hyaline in 3% KOH to light yellowish in
Melzer's solution. Subhymenium: a narrow layer of thin-
walled, short, filamentous, hyphae 1.7-2.5 um diam, hyaline
in 3% KOH, with scattered clamp connections. Basidia: 19-35
X 6-8 um, clavate, laginiform, to clavate-rostrate, thin-
walled, 4-spored, with infrequent 1 and 2 spored basidia,
hyaline in 3%KOH and Melzer's solution. Partial veil: of
filamentous, thin-walled, hyaline, hyphae 2.5-6 (-8.5) um
diam infrequently branched at right angles or Y shaped with
frequent clamp connections. Universal veil not present.
Basidiospores: 6-7.5 X 5-6 um diam, subglobose, globose,
thin-walled with low, brown, warts and a small to
indistinct, hyaline, apiculus, yellow-brown in 3% KOH and
Melzer's solution. Spore deposit brown.
Habitat and distribution: terrestrial, common, gregarious,
some subcaespitose, in mixed pine and hardwood forest or in
the beech, magnolia-loblolly pine mixed forest, often near
baygall communities. In Southeastern Texas. Fruiting in
spring from May until June and again in October.
Material examined. United States. Texas: Orange Co.:Vidor,
23 May 1977, Lewis 974% Vidor, 5 May 1979, Lewis, 1719;
Vidor, 16 July 1989, Lewis 4297; Hardin Co., Lumberton, 4
Oct. 519797) Lewis 20357) Vidor) (28) May, 1989s. bewrs 74250;
Hardin Co., Lance Rosier Unit, Big Thicket National
Preserve, 13 May 1990, Lewis 4370 (Holotype); Orange Co.,
Sam Houston Nat. Forest, 2 Nov. 1979, O.K. Miller 18684.
Austria. Tirol: Vikartal, Cortinarius nothosaniosus, 9
Sept. 1951, M. Moser Nr.51/120 (holotype) (I). Sweden.
Femsjo: ost Trollgol, Cortinarius limonius, 8 Sept. 1974, M.
Moser, Nr.74/328 (1). Femsjo: Aborrasjo, Cortinarius
Fimoneus,;) 4 Augeuloy2, “MayMoser;, Nr wiz /1oo (iy
Observations: In the baygall communities Cortinarius lewsii
is often gregarious and numerous. The bright, orange-yellow,
dry, pileus, yellow flesh, (Metzler et al 1992) and small
subglobose to globose spores, and a pileipellis of
interwoven hyphae are a distinctive set of characteristics
which would place this species in the Subgenus Leprocybe,.
The section Limonei has species with vivid yellow to yellow
brown pileus coloration and globose, subglobose, to broadly
elliptical, warted spores. It appears to be close to C.
nothosaniosus Moser which is found in "versauerter
Fischtenwald (Picea abies) mit Vaccinium myrtillus"” in
Europe, (Moser, 1966). However, C. nothosaniosus has a more
acute umbo as illustrated by Moser (1966). In addition, the
pileus is smaller (up to 2.5cm broad), the lamellae
subdistant and the stipe thin and small (1.5-4cm X 2-4mm
thick). Buttons are red brown with a yellowish margin which
is fibrillose from the veil. Microscopic features are
465
Similar to those of C. lewisii. The basidioles of C.
lewisii are variable in shape as described above and
probably result from the maturation of 1, 2 and 4 spored
basidia.and distinctive clusters of hyphae are present in
the pileipellis with cells 8-30 pum diam. Cortinarius
Limonius (Fr. Jex) Fro) Fr. is) similar’ but hasian'|"foxy, red"
to "orange red" pileus, is more robust than C. lewisii, and
has a clavate stipe. It is a European species also found
associated with spruce and pine (Moser & Julich 1986,
Moser), 1933)",
Species that produce orellanin are found in the
Subgenus Leprocybe. Testing for orellanin by Poder and Moser
(1989) have revealed that the toxic species are in the
Section Orellani. When tested neither C. nothosaniosus nor
Cc. limonius, in section Limonei, contain orellanin. It is
logical to conclude that C. lewisii also is unlikely to
containmene toxin:
ACKNOWLEDGEMENTS
I thank David Lewis for notes and collections and the
Texas Mycological Society for their hospitality. Susan and
Van Metzler provided support and help. Dr Harold Burdsall
and Dr. Joe Ammirati provided much appreciated comments on
the manuscript. Logistical support was provided by the
Virginia Museum of Natural History.
LITERATURE CITED
Fries, E. 1821 Systema Mycologica I: 213.
Metzler, S., V. Metzler, & O.K. Miller. 1992. Texas
Mushrooms. Univ. of Texas Press, Austin, Texas. 350p.
Moser, M. 1966. Einige interessante Pilzfunde aus dem Gebiet
von Gotschuchen. Sonderdruck aus Carinthia II, Mitteilungen
des Naturwissenschaftlichen Vercines fur Karnten, 76, bzw.
DSO e390.
Moser, M. 1983. Keys to Agarics and Boleti. Gustav Fischer
Verlag, Stuttgart, ‘Germany 535 ).
Moser, M. & Julich 1986. Farbatlas der Basidiomycetes. III
Agaricales. Cortinarius 3. Gustav Fischer Verlag
Kornerup, A. and J.H. Wanscher. 1966. Methuen handbook of
SoLlour.ndled. .Metnuenta.CO.,) LUtLa.e) London,
Poder, R. und M. Moser. 1989. Eine einfache, empfindliche
methode zum makrochemischen nachweis von orellanin.
Mycologia Helvetica 3: 283-290.
Ridgway, R. 1912. Color standards and color nomenclature.
Washington D.C. 44p., I-LIII Pl.
466
Watson, G. 1979. Big thicket Plant Ecology. 2nd Ed. Big
Thicket Museum, Saratoga, Texas 10 p.
Fig. 4. Cortinarius lewisii xX Le
MY COTAXON
Volume XLVII, pp. 467-474 April-June 1993
BOOK REVIEWS
L. M. Kohn, Book Review Editor
Mycota Lithuaniae Volume 5 (1): Rudieciai (Uredinales), by A.
Minkevicius & M. Ignataviciute. Hardcover, 223 pp., 240 x 170 mm,
1991. Published by the Lithuanian Academy of Sciences. Requests
must be sent to the Institute of Botany (Zaliyju ezery 47, Vilnius, 2021,
Lithuania), fax 01 22 359950.
This represents the first in a series of of floristic volumes which
will eventually cover all major groups of fungi known to occur in
Lithuania. The present fifth volume was published out of sequence due
to the advanced age of its first author, Professor Minkevicius, who has
studied the Basidiomycota at the Lithuanian Academy of Sciences over
the last 50 years. Although this first volume contains only line
drawings, future plans for the series include color photo illustrations of
macrofungi as well as detailed taxonomic descriptions of new taxa.
The current volume includes keys and descriptions to about 170
species of rust fungi from Lithuania, with data on their distribution and
host biogeography. The introductory chapters also include interesting
discussion on rust life history, and particularly the apparent rarity of
certain life cycle stages (mostly aecial) in many species owing in part to
climatic conditions in this Baltic country.
The Mycota Lithuania project was conceived and its first volume
published while Lithuania was still a Soviet state. Although economic
problems of a free Lithuanian state will certainly delay publication of
the remaining volumes already underway, the eventuality of a complete
documented mycota for Lithuania will represent a valuable contribution
for understanding the biogeography of European fungi. Although this
first volume is written in Lithuanian, it contains excellent summaries in
both Russian and English, and most of its technical information
concerning taxonomic characters and distributions should still be
readable and of interest to other professional mycologists working with
these fungi. -R. Vilgalys, Department of Botany, Duke University,
Durham , North Carolina 27708 U.S.A.
Foliicolous Lichens - A Contribution to the Knowledge of the
Lichen Flora of Costa Rica, Central America, by Robert Liicking.
Beihefte zur Nova Hedwigia, Heft 104. VII: 1-179 + 64 figs. Softcover,
179 pp., 240 x 170, 1992. J. Cramer in der Gebriider Borntraeger
Verlagsbuchhandlung, Johannesstr. 3 A, D-7000 Stuttgart 1, Germany,
fax (0711) 625001. ISBN 3-443-51026-4. DM 170.--.
The principal component of this publication are keys to all (c. 230)
obligately foliicolous lichen species known from Costa Rica. Most of
these have been found there for the first time by the author during his
468
extensive investigations, including several new species published in a
separate paper (Nova Hedwigia 52: 267-304). Added are very useful,
good quality, black and white photographs of most species, and many
line drawings of spores, asci, paraphyses, etc. Costarican localities and
world distributions are provided for each species, and for most species,
key, discriminative characteristics are pointed out. The short general
part contains a list of collection sites and a glossary.
This book transforms Costa Rica into the country with the second
largest known foliicolous lichen flora in the world. The size of this
flora probably results from the varied topography of the country. It
certainly also results from the poor knowledge of the lichen flora in
other tropical countries.
Facultatively foliicolous lichens are poorly represented in the book.
These may be recognized because they tend to also grow on adjacent |
twigs. They are by no means rare and are sometimes really
conspicuous. Either Lticking happened not to come across them, or he
omitted them, not an unusual practice. Further, a number of common
neotropical taxa, like Mazosia tumidula, are missing; evidently these
have not yet been found, or do not occur, in Costa Rica. For evident
reasons the data on world distribution should be considered as
provisional. The ecological rather than taxonomic background of the
author is reflected in some problematical taxonomic practices. The
introduction of "ad interim" names seems undesirable because they may
come into wider use through the keys and lead to nomenclatoral
confusion. Further, the "isotypes" do not completely reflect current use
because they bear different numbers; however, they do not really
deviate from the common practice of including in a single collection a
number of individuals growing together. In this treatment, individuals
which grew on different nearby shrubs have been given different
numbers.
The main significance of this book is as an identification tool. It
forms a very valuable addition to Santesson's monographic treatment of
1952, the main source of information for the group, because it treats
numerous species which have been discovered since 1952, and follows
the newest taxonomic rearrangements. Moreover, the numerous
illustrations are a welcome complement to Santesson's sparsely
illustrated book. The keys contain most of the important characters,
making descriptions largely superfluous, and the illustrations capture
subtle characters which are difficult to aptly convey in words.
H. Sipman, Botanischer Garden & Botanisches Museum, Konigin-
Luise-Stragpe 6-8, D-1000 Berlin 33, Germany.
469
The Polypores of China, by Zhao Ji-Ding & Zhang Xiao-Qing.
Bibliotheca Mycologica Band 145. Softcover, 524 pp., 225 x 140 mm,
1992. J. Cramer in der Gebriider Borntraeger Verlagsbuchhandlung,
Johannesstr. 3 A, D-7000 Stuttgart 1, Germany, fax (0711) 625001.
ISBN 3-443-59046-2. DM 190.--.
This flora, written in English, treats 351 species. One new
combination is proposed, Phellinus quercinus (Zhao) Zhao, and no new
species are described. A short introduction is followed by a list of the
Chinese polypores. There is a key to the 76 genera and keys to species
appear under the treatment of each genus. The generic concepts in most
cases follow Gilbertson and Ryvarden's North American Polypores,
1986-1987. The polypores in the major works on Chinese fungi by Tai
(Sylloge Fungorum Sinicorum, 1979) and Teng (Fungi from China,
1963) have been incorporated.
The species’ descriptions are original having been prepared from
_ Specimens studied by the authors. They are detailed and follow a
consistent format. The principal microscopic features of 318 species
are illustrated by line drawings. Each species treatment includes data
on habitat, specimens examined, and distribution. The sources for the
notes on habitat were probably comments accompanying the specimens,
but some comments seem to be from the literature. The distribution
simply lists some countries where the species has been reported. A
surprisingly large number of the species, e.g., 53 of the first 100, also
occur in the northern United States and Canada.
Those using this text to identify collections should be aware of a
few potential problems. First, in the key to the genera each generic
name appears only once. Thus, a genus having some species with and
others without clamp connections, appears only once under one key
choice, e.g., with clamp connections, and it will be difficult to correctly
key-out those species lacking clamp connections. Second, Meruliporia
violacea with hyaline, thin-walled spores is obviously misplaced. The
prevailing concept of Meruliporia is of a genus with distinctly yellow-
brown spores. Third, in some drawings the spore shapes differ
somewhat from North American specimens, i.e., the spores are overly
attenuated at the apiculus end. Fourth, the size of the spores in the
drawings sometimes differs from the sizes given in the text, e.g., for
Dichomitus squalens the text says spores 7-8 Um long but the drawing
has spores 6-9.5 um long. Finally, the spore ornamentation in
Bondarzewia montana is correctly described as "short, irregularly
arranged ridges," but the illustration shows five spores with conical to
hemispherical warts.
This is an impressive volume. It is well-organized, detailed and
readable, except for some instances where the choice of words is
confusing. It will be an important reference in the identification of
specimens from Asia. Students of the polypores will find this a useful
volume, as will biogeographers, forest pathologists and plant quarantine
scientists. J. Ginns, Centre for Land and Biological Resources
Research, Ottawa, Canada.
470
Pilze auf Pilzfruchtkorpern. Untersuchungen zur Okologie,
Systematik und Chemie, by Wolfgang Helfer. Libri Botanici 1:1-157
+ 8 color figs. Softcover, 240 x 170 mm, 1991. [HW-Verlag, Bert-
Brecht-StraBe. 18, D-8057 Eching, Germany, fax 089-3192257. ISBN
3-9802732-2-9. DM 88.--, US $55.--.
This initial volume of a new series concerns the fungi that occur on
other fungi, sometimes referred to as the fungicolous fungi or
fungicoles. The book consists of two basic sections. One section
presents a taxonomic account of the 140 species of Bavarian
fungicolous fungi encountered in the study including seven new
species. Although primarily Ascomycetes and Fungi Imperfecti, there
are 9 species in the Zygomycetes and 13 Basidiomycetes. Fungi
occurring on myxomycetes are listed separately. Many of these species
occur widely in temperate regions. Descriptions are included for only
the newly described species with taxonomic literature for each species
abundantly cited. A few taxonomic inconsistencies exist, for example,
Nectria vilior Starb. (as N. violor) is listed as a synonym of N.
berkeleyana on p. 52 while on p. 47 it is not and should not be in my
estimation. But for the most part the taxonomy is accurate and up-to-
date. This section combined with the list of substrates, a key to genera,
and a key to the known or suspected anamorphs of Hypomyces make
this book an extremely useful synopsis of fungicolous fungi. The key
to anamorphs of Hypomyces is especially useful given that many
Hypomyces are encountered only rarely as the teleomorph but may have
one or two distinctive anamorphs. Results of a chemotaxonomic
analysis of anamorphs of Hypomyces bring into question current
concepts of these anamorph genera. The second section presents
experimental data on factors that influence infection and host specificity
of Hypomyces chrysospermus, a fungicolous species thought to be
restricted to members of the Boletales. In one series of experiments,
extracts of various host fungi are shown to affect the germination rate of
aleurioconidia and phialoconidia differentially. Other experiments
involve the effects of extracts of Paxillus involutus on aleurioconidia
under various conditions. Finally, an attempt was made to isolate and
identify the factor that stimulates germination of aleurioconidia.
Although unsuccessful, this research is unique and fascinating with
implications for related fields, for example, circumscription of the
Boletales. A. Y. Rossman, USDA-ARS Systematic Botany and Mycology
Laboratory, Beltsville, Maryland 20705-2350 U.S. A.
471
Ektomykorrhizien an Fagus sylvatica: Charakterisierung und
Identifizierung, okologische Kennzeichnung und unsterile
Kultivierung, by Felix Brand. Libri Botanici 2: 1-229. Softcover, 240
x 170 mm, 1991. IHW-Verlag, Bert-Brecht Strafe 18, D-8057 Eching,
Germany, fax 089-3192257. ISBN 3-9802732-4-5.
DM 105.--, U.S. $ 65.--.
Since the mid 1980's, Reinhard Agerer and his students have
published several studies on the anatomy and morphology of various
ectomycorrhizae in their well known series of publications entitled
"Studies on Ectomycorrhizae" as well as in the Colour Atlas of
Ectomycorrhizae. Now, a compilation of studies by Felix Brand is
available on the ectomycorrhize of European beech.
Keys are provided for the identification of 23 distinct beech
ectomycorrhizae along with their fungal symbionts. A detailed
description including line drawings and black and white photographs of
the morphology and anatomy is provided for each ectomycorrhiza in
question. In addition, the manual contains sections on how to prepare
and examine field-collected ectomycorrhizae and how to interpret the
descriptions provided in the text. Detailed line drawings aid in this
respect. Comparisons are made with reports of ectomycorrhizae on
beech by other authors. A whole chapter provides detailed information
on the edaphic factors and general ecology of the ectomycorrhizal fungi
associated with beech, while another deals with the synthesis of beech
ectomycorrhizae under laboratory conditions.
The specialized scope of the book may limit its usefulness.
However, the detailed information presented cannot be overlooked by
any researcher working on the anatomy and morphology of
ectomycorrhizae found on other tree species. L. J. Hutchison,
Department of Forest Science, University of Alberta, Edmonton , Alta.,
Canada.
Book Review Editor's note: MYCOTAXON has received three
additional volumes in this series. Libri Botanici 3: 1-215; Allgemeine
Taxonomie und Chorologie der Pflanzen, by Werner Rothmaler, a 1992
reprint of a work originally published in 1955 (ISBN 3-9802732-5-3,
DM 36.--). Libri Botanici 4: 1-115 + map; Die Vegetationsentwichlung
in NaBbaggerungen der bayerischen Donauebene, by Andreas Otto,
1992 (ISBN 3-9802732-7-X, DM 68.--). Libri Botanici 5:1-288 + 16
color photographs, Pilzflora von Bayreuth und Umgebung, by
Wolfgang Beyer, 1992 (ISBN 3-9803083-1-6, DM 88.--). Note that the
Pilzflora includes many ascomycetes (with some nice color photos) as
well as basidiomycetes; Red List status is indicated for some entries.
472
An Illustrated Manual on Identification of some Seed-borne
Aspergilli, Fusaria, Penicillia and their Mycotoxins, by K. Singh, J.
C. Frisvad, U. Thrane, and S. B. Mathur. Softcover, 133 pp., 296 x 210
mm, 1991. Danish Government Institute of Seed Pathology for
Developing Countries, P.O. Box 34, Ryvangs Allé 78, DK-2900
Hellerup, Copenhagen, Denmark. ISBN 87-7026-3175. Price outside
Denmark is DKK 220.-- (excl. VAT) + postage.
This manual was written to provide a means of identifying
seedborme species of Aspergillus, Fusarium, and Penicillium,
with an emphasis on mycotoxins and other secondary metabolites
as identification criteria. The book begins with information
on techniques for isolating these fungi into pure culture, for growing
them under standard conditions for morphological observations, and for
detecting mycotoxins in culture. A compilation of informationon
physical characteristics of mycotoxins is provided, e.g., UV spectra,
TLC data and HPLC retention indices. Descriptions illustrated with
mostly color plates and line drawings are provided for 15 aspergilli, 9
fusaria, and 18 penicillia. A synoptic key is provided for species treated
in each genus. The final chapter details methods for detecting and
identifying mycotoxins in seeds. The book ends with a glossary of over
40 terms and a bibliography arranged by subject, with comments on
some of the works cited. The authors state that they have followed the
modern taxonomic and nomenclatural approach outlined in Samson and
Pitt, Eds. (Plenum Press. 1990).
I find two main limitations to this manual. First, the color
terminology is general and lacks reference to a standard. For example,
“creamish yellow" describes the color of the reverse of a number of
colonies that are shown in the plates with colors that differ significantly
from each other. Additionally, for some species the color seen in the
plate does not correlate with the color indicated in the description.
Second, measurement data of microscopic structures are not provided,
presumably because the information is available elsewhere and because
this book emphasizes the use of mycotoxins for identification. This
emphasis assumes a lack of variation in mycotoxin production within a
species. It also requires that additional references be used to identify an
isolate with certainty. For these reasons, the book is less useful as a
lone identification manual, especially for users not already familiar with
these organisms.
Despite these limitations, this book successfully addresses the
needs of a user group that must identify species in this difficult group.
The color photomicrographs are generally of high quality. The most
valuable aspect of the book is the fact that it is a concise source of
identification information for these three genera, especially regarding
mycotoxin detection techniques. Users will find this book a worthwhile
addition to their collection of identification tools. M. E. Palm,
USDA/APHIS, Systematic Botany and Mycology Laboratory, Beltsville,
Maryland 20705-2350 U.S.A.
473
Pleurotus unter Ste}. Okophysiologische Untersuchungen zu
Wasserhaushalt und Sporulation, by Angelika Achhammer.
Bibliotheca Mycologica Band 141. Softcover, 206 pp., 225 x 140 mm,
1992. J. Cramer in der Gebriider Borntraeger Verlagsbuchhandlung,
Johannesstr. 3 A, D-7000 Stuttgart, Germany, fax (0711) 625001.
ISBN 3-443-59042-X. DM 130.--.
This doctoral thesis reports on the role of water relations in
vegetative growth, fruiting, and sporulation of several isolates of
Pleurotus ostreatus and one of P. pulmonarius. Emphasis is on the
response to water stress induced either directly or indirectly. Long term
and daily sporulation patterns were determined in the field and under
various combinations of controlled temperature, humidity and duration
of daylight and darkness.
Water loss, uptake, and transport were studied with detached
fruitbodies taking into account differences among major fruitbody
regions. Although sporulation continued despite considerable
dehydration, even very high relative humidity could not revive dry
fruitbodies. Liquid water was needed for resumption of sporulation,
and to be effective had to be supplied through the cap because water
movement through the stipe was generally too slow. Total water and
osmotic potentials of fruitbodies were determined, and the effects of
drying and freezing were studied by scanning electron microscopy of
different fruitbody regions, especially the hymenium. Vitality was
tested with fluorescein diacetate. Tolerance of osmotic stress was
studied for mycelial growth, fruiting, sporulation and spore germination
in cultures grown on media with added osmotica (NaCl and for some
tests glycerol). The content of trehalose and several polyols was
determined in fruitbodies subjected to osmotic stress and freezing
because these compounds may protect against water stress.
The book will interest researchers studying the physiology of water
relations and ecology of Hymenomycetes. It will also interest
Pleurotus growers and forest pathologists. There is a three-page
summary in English and an extensive bibliography. Few typographical
errors were noted. Unfortunately, readability suffers from the fact that
the results are subdivided into many sections each with introductory
remarks, literature review and a lengthy discussion. In consequence it
is difficult to focus on the results and to get a clear overview of the
material. The problem is increased by the large number of
abbreviations and the excessively complex numbering of the figures.
H. E. Gruen, Department of Biology, University of Saskatchewan,
Saskatoon, Sask., Canada.
Texas Mushrooms: A Field Guide, by Susan Metzler and Van
Metzler; Scientific Advisor, Orson K. Miller, Jr. Hard or Softcover,
304 pp. with 249 color photographs, 5.75 x 8.25 inches, 1992.
University of Texas Press, P.O Box 7819, Austin, Texas 78713-7819.
ISBN (cloth hardcover) 0-292-75125-7, U.S $39.--; ISBN (paper
softcover) 0-292-75126-5, U.S. $17.95.
474
The authors describe their book, Texas Mushrooms, as a pictorial
guide for beginners to common Texas mushrooms and fungi. It
contains 222 major entries, each with at least one color photograph.
Nearly 40% of the descriptive part of this book covers just the boletes,
Amanita, Lactarius, and Russula. Several of the taxa included here
cannot be found illustrated in other field guides. Many of the
photographs are quite striking and the book is well bound and printed
on good paper. Also, this is the first mushroom field guide of Texas. In
fact, despite its short-comings, some particular to this book, some
common to the field guide genre, Texas Mushrooms is a welcome
addition to the growing literature of regional guides, and can give
residents and non-residents alike a sense of this particular
geographically defined mycoflora that is unavailable from any of the
national field guides. |
The book is divided into five parts: and introduction to mushroom
hunting in Texas; how to use the book to identify mushrooms
(illustrations, charts, etc.); a section on mushroom toxins by Robert
Harvey, M.D.; a section on cooking and eating wild mushrooms (with
10 recipes); and the bulk of the book, some 271 pages, with text and
accompanying photographs on the same page. There is a short list of
books for further study and an index. There is also an appendix of
spore data, despite the authors’ stated preference for choosing "species
that can be clearly distinguished by field characteristics," and their
statement that "when (microscopic) examination is the only way to
distinguish between closely related species, (they) provided the name of
the most commonly found form" (p. 3).
Because field guides are part of taxonomic literature but are not
scientific literature (usually no voucher collections are cited, etc.), the
most pressing question becomes: what is an acceptable level of
imprecision, and for whom? Texas Mushrooms lacks authors'
citations, specific references to technical descriptions with each major
entry, and voucher collections. Nowhere does it state that all the
photographs were even taken in Texas, and, despite this guide's reliance
on the photography for species determinations, some of the photos are
not diagnostic, or they are murky, or badly off-color, or misdetermined.
The species descriptions are less than adequate, and they are wrong in
detail for some entries and neglect essential field characters of others.
In fact, many species descriptions seem intended to cover whole
complexes or even genera rather than species. There are also examples
of the same species represented by different entries (different names
and photographs). The section (and scattered notes) on mushroom
poisoning is badly misleading and in error in several instances. G.
Lincoff, The New York Botanical Garden, Bronx, New York, U.S.A.
MY COTAXON
Volume XLVI, pp. 475- 479 April-June 1993
NOTICE: A NEW COMPUTER PROGRAM
FOR THE IDENTIFICATION OF LICHEN SUBSTANCES
ESTHER MIETZSCH
Institut fiir Physiologie, Abt. Biokybernetik, Ruhr-Universitat Bochum, D-4630 Bochum,
Germany
H.THORSTEN LUMBSCH
Botanisches Institut, Universitat Essen, Postfach 103 764, D-4300 Essen 1, Germany
JOHN A. ELIX
Department of Chemistry, The Faculties, Australian National University, GPO Box 4,
Canberra, ACT 2601, Australia
Abstract: A program for the identification of lichen substances based on
TLC RF values, HPLC RI-values, colours of developed spots under visible
and UV light, as well as the results of lichen spot tests has been developed
for Microsoft WINDOWS. The program is based on a similar program for
Apple Macintosh computers. It accepts RF values from six standard
solvents, a RI-value from a gradient HPLC with reverse phase column and a
range of possible colours for an unlimited number of TLC spots and will
generate answers within a user defined error range. A list of synonyms of
lichen substances is included as well as an RF and RI-value calculator.
Introduction
Chemical investigations of lichens form an integral part of all serious taxonomic
studies on lichen-forming fungi today. About 550 lichen substances are known, most
of them specific to lichens. The significance of chemical characters in the taxonomy
and systematics of lichenized fungi has often been discussed (W. CULBERSON 1970,
1986, C. CULBERSON 1986, CULBERSON & CULBERSON 1970, HAWKSWORTH 1976,
BRODO 1978, 1986, EGAN 1986). However, even the more experienced lichenologist
476
encouters TLC spots or HPLC peaks that are unfamiliar and difficult to identify. Lists
of RF-values in standardized TLC solvent systems (CULBERSON & KRISTINSSON 1970,
CULBERSON 1972, CULBERSON, & AMMANN 1979, ELIx et al. 1987b) and RI-values
of gradient elution HPLC (FEIGE et al. 1993) have been published. These lists,
however, cannot be kept current since new substances are continuing to be discovered
and a large bulk of scattered literature exists on the RF-values of recently described
substances. Associated information such as the colour of the spots, mass spectra data
etc. is not available in lists, but is scattered within the literature. In an effort to make
the information more readily accessible, and to keep such a library of information
current as more and more lichen metabolites are identified and characterised, a data
bank suitable for storing such information was developed by ELIX et al. (1987a, 1988)
for APPLE MACINTOSH computers.
Discussion
In the past it has been very difficult to produce a user-friendly menu environment in
the Dos system. However, with the improvement in personal computers and the
introduction of the WINDOWS system this situation has changed. Moreover powerful
computer equipment is now available at reasonable prices.
The design of the menu environment and windows of the new program resembles that
of the MACTABOLITES program, but three further options were added (MIETZSCH et al.
1992). These are a "Calculate" menu to calculate RF- and RI-values, "Synonyms",
which gives a list of synonyms and the appropriate names in current use and a "Help"
menu. The substance classes and the HPLC RI-values are additional features of
WINTABOLITES. The program makes full use of the user friendliness of the WINDOWS
system as illustrated in Figure 1. The program accepts RF-data from six standard
solvents, an RI-value for a gradient HPLC with reverse phase column and a range of
possible colours for an unlimited number of TLC spots. A list of possible answers is
generated within a user defined error range of RF/RI-values and colours.
The data set operates primarily on RF-values obtained by TLC on silica gel in any
number of six standard solvent systems as well as the gradient HPLC system. Thus
listed along with the name of each lichen substance is the following information:
1. The TLC RF-values of the compound in six standard solvent systems:
A, toluene/ dioxane/ acetic acid (180 : 45 : 5)
B, hexane/ diethyl ether/ formic acid (130 : 80 : 20)
B’, hexane/ methy!] tert-butyl ether/ formic acid (140 : 72 : 18)
C, toluene/ acetic acid (170 : 30)
E, cyclohexane/ ethyl acetate (75 : 25)
G, toluene/ ethyl acetate/ formic acid (139 : 83 : 8)
2. The HPLC RI-value from the gradient system described by FEIGE et al. (1993) for
phenolic compounds.
3. Whether the TLC spot is coloured in visible (natural) light and/or detectable in
477
short wavelength ultraviolet light.
4. The colours of the TLC spot in both visible (natural) and long wave length ultra-
violet light after spraying with H2SO4 and charring.
5. The colour of the TLC spot after spraying with Archers solution.
6. The results of medullary spot tests with K, C, KC, PD (when a compound gives a K
+ or C + reaction then KC has been omitted unless a different colour is produced
by KC).
7. Three major peaks in the mass spectrum plus molecular ion (if observed).
8. The substance class.
=z Wintabolites
File Edit Data Search Calculate Synonyms Help
| 6-O-Methyithiophanic BIOSYNTHETICALLY RELATED COMPOUNDS
65 x 71 60
2.5, 7-Trichloro-3-O-methylnorlichexanthone
72 x 55 66 3-0-Methylasemone
lisohypertatolic acid 5,7-Dichloro-3-o-methyinorlichexanthone
lsoarthothelin
Thiophanic acid
Thuringione
12,4-Dichlorolichexant
78 x «676 O75
iHyperlatolic acid
53 x 74 52
2,5, ?-Trichlorolichexanthone V: + UV: +
87 x 74 85 58 x 54
Fig. 1: WINTABOLITES - Compounds Biosynthetically Related to 3-O-Methylthio-
phanic acid; Screen Display.
9. Up to nine biosynthetically related compounds.
10. Notes, a one hundred character long message containing additional information
regarding the characteristics of the particular substance.
However, it is not necessary to have all such information about an unknown spot - the
search program will operate on as little information as a single RF or RI-value.
Qualifying information simply reduces the number of possible answers generated.
478
The program was written in TURBO PASCAL for WINDOWS (Borland) using object-
oriented programming techniques. The size of the source code is 66 KB. As all data
are kept in the main memory by using dynamic data structures during the running of
the program, a very high speed of retrieval is achieved. After the program has started,
the data are loaded from the data file and all records are sorted into a binary tree
which ensures quick access to the data (WIRTH 1983). At the same time seven arrays
are build up, there being a separate array for each solvent system and one for the
HPLC. The arrays are indexed by the RF-value or the RI-value of the compounds and
the elements of the arrays each contain a dynamic list of pointers to all the records
which have an RF or RI of that value for that solvent. When a search is started all the
lists in the array of the first solvent system from RF - error index to RF + error index
of the first spot are checked and all records, where the additional information given in
agreement, are regarded as possible answers. The number of data sets as well as the
number of spots to be examined is unlimited.
WINTABOLITES is a WINDOWS application. Its use requires WINDOWS 3.0 or 3.1. the
computer must have a 80286 or higher (i.e. 80386 or 80486) CPU and at least 2 MB
RAM. A VGA graphic card and monitor is recommended.
Availibility of the program: The program and an illustrated manual are available from
the German authors for a price of US $ 149. Private persons not attached to an
institution may ask for a reduced price of US $ 79. Updates will be distributed to the
WINTABOLITES users yearly for DM 10 to 15,- if they wish to receive them.
We wish to thank Dr. Siegfried Huneck (Halle) who kindly reviewed this manuscript.
References
BRODO, I.M. (1978): Changing concepts regarding chemical diversity in lichens.
Lichenologist 10: 1-11.
BRODO, I.M. (1986): Interpreting chemical variation in lichens for systematic purposes.
Bryologist 89: 132-138.
CULBERSON, C.F. (1972): Improved conditions and new data for the identification of lichen
products by a standardized thin-layer chromatographic method. J. Chromatogr. 72:
113-125,
CULBERSON, C.F. (1986): Biogenetic relationships of the lichen substances in the
framework of systematics. Bryologist 89: 91-98.
CULBERSON, C.F. & K. AMMANN (1979): Standardmethode zur Diinnschichtchromato-
graphie von Flechtensubstanzen. Herzogia 5: 1-24.
CULBERSON, C.F. & H.D. KRISTINSSON (1970): A standardized method for the identifi-
cation of lichen products. J. Chromatogr. 46: 85-93.
CULBERSON, W.L. (1970): Chemosystematics and ecology of lichen-forming fungi. Ann.
Rev. Ecol. Syst. 1: 153-170.
CULBERSON, W.L. (1986): Chemistry and sibling speciation in the lichen-forming fungi:
ecological and biological considerations. Bryologist 89: 123-131.
CULBERSON, W.L. & C.F. CULBERSON (1970): A phylogenetic view of the chemical
479
evolution in the lichens. Bryologist 73: 1-31.
EGAN, R.S. (1986): Correlations and non-correlations of chemical variation patterns with
lichen morphology and geography. Bryologist 89: 99-110.
ELIx, J. A., JOHNSTON, J., and PARKER, J. L. (1987a): Mactabolites. Users Manual. 2nd ed.
Canberra.
ELIx, J.A., J. JOHNSTON & J.L. PARKER (1987b): A catalogue of standardized thin layer
chromatographic data and biosynthetic relationships for lichen substances. Australian
National University, Canberra.
ELIx, J.A., J. JOHNSTON & J.L. PARKER (1988): A computer program for the rapid
identification of lichen substances. Mycotaxon 31: 89-99.
FEIGE, G.B., LUMBSCH, H.T., HUNECK, S. & ELIX, J.A. (1992): A method for the
identification of aromatic lichen substances by gradient high performance liquid
chromatography. J. Chromatogr., in prep.
HAWKSWORTH, D.L. (1976): Lichen Chemotaxonomy. S. 139-184. In: D.H. BROWN, D.L.
HAWKSWORTH & R.H. BAILEY (Hrsg.): "Lichenology: Progress and Problems", Aca-
demic Press, London.
MIETZSCH, E., H.T. LUMBSCH & J.A. ELIX (1992): Wintabolites (Mactabolites for
Windows) Users Manual. Universitat Essen.
WIRTH, N. (1983): Algorithmen und Datenstrukturen. 3rd. edn. Stuttgart, Teubner Verlag.
ie
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481
Author Index to Volume Forty-seven
Abdullah, N. see Ho, Y. W., D.J.S. Barr, N. Abdullah, and S. Jalaludin
see Ho, Barr, Abdullah, Jalaludin and Kudo
Alvarez, Isabel F. Javier Parladé, James M. Trappe and Michael A. Castellano
Hypogeous mycorrhizal fungi of Spain. 201-217
Bakalova, Ganka G. see Sameva and Bakalova
Banerjee , Partha and Walter J. Sunberg. Three new species and a new variety of
Pluteus from the United States. 389-394
Barr, D. J.S. see Ho, Y. W., D.J.S. Barr, N. Abdullah, and S. Jalaludin
see Ho, Barr, Abdullah, Jalaludin and Kudo
Blackwell , Will H. and Martha J. Powell. Taxonomic history of the oomycete genus
Thraustotheca. 183-192
Boqueras , Montserrat and Paul Diederich. New or interesting lichenicolous fungi.
3. Karsteniomyces llimonae sp. nov. and Sclerococcum SeruSiauxii sp. nov.
(Deuteromycotina). 425-431
Castellano, Michael A. see Alvarez, Parladé, Trappe and Castellano
Corlett, Michael Taxonomic studies in the genus Mycosphaerella. 2. Notes on some
additional species occurring on Brassicaceae. 131-146
Diederich, Paul see Boqueras and Diederich
Eicker, Albert , J. V. Van Greuning and Derek A. Reid. Amanita reidii - a new species
from South Africa. 433-437
Elix, John A. New species in the lichen family Parmeliaceae (Ascomycotina) from
Australia. 101-129
and Debra A. Venables. 4-O-Methyllividic acid, a new lichen depsidone. 275-281
Frizzi, G. see Pacioni, Frizzi, Miranda and Visca
Galan, R. Moreno and Galan
Gomez-Bolea, A. see Muntafiola-Cvetkovic and Gémez-Bolea
Grosse-Brauckmann., Helga see Hjorstam and Grosse-Brauckmann
Guzman, Gast6n see Ryvarden and Guzman
Haines, John see Olsen, Haines and Sivertsen
Halling, Roy E. Two new Crinipellinae (Tricholomataceae: Marasmiae) from South
America. 379-385
Hess, W. W. see Strobel, Stierle, Stierle and Hess
Hjortstam, Kurt and Leif Ryvarden. Two new South American species of
Corticiaceae (Basidiomycetes) with amyloid spores. 81-85
and Helga Grosse-Brauckmann. Two new species of Cristinia (Basidiomycotina,
~ Aphyllophorales) and a survey of the genus. 405-410
Ho, Y. W., D. J. S. Barr, N. Abdullah, and S. Jalaludin. Anaeromyces, and earlier
name for Ruminomyces. 283-284
and H. Kudo. A new species of Piromyces from the rumen
of deer in Malaysia. 285-293
_____and§&. C. Jong. Phytophthora hibernalis and P. syringae. 439-460
Ialongo, Marco T. Biostatistical characterization of the genus Oidium. 193-199
Jalaludin, S. see Ho, Barr, Abdullah, and Jalaludin
see Ho, Barr, Abdullah, Jalaludin and Kudo
Jayachandra , J. A. see Muthumary, Jaychandra and Preetha
Jong , Shung-Chang and Candace McManus. Computer coding of strain features of the
genus Rhizopus.. 161-176
see Ho and Jong
Ju, Y..-M. Felipe San Martin Gonzalez and Jack D. Rogers. Three xylariaceous fungi
with scolecosporous conidia. 219-228
Kohn, L. M. Book Reviews. 467-473
482
Kudo, H. see Ho, Barr, Abdullah, Jalaludin and Kudo
Kuthubutheen, A. J. see McKenzie and Kuthubutheen
McKenzie, E. H. C. and A. J. Kuthubutheen. Dematiaceous Hyphomycetes on
Freycinetia (Pandanaceae). 4. Cryptophiale. 87-92
McManus, Candace see Jong and McManus
Miller, Orson K. A new Cortinarius from Southeastern Texas. 461-466
Miranda, M. see Pacioni, Frizzi, Miranda and Visca
Moreno, G. and R. Galan Hydnangium pila Pat., an older name for Martellia
mediterranea Moreno, Galan & Montecchi. 157- 159
Morgan-Jones, G. see Phelps and Morgan-Jones
Muntanola-Cvetkovic, M. and A. Gémez-Bolea. Chaetospermum chaetosporum
(Coelomycetes). First record from the Iberian Peninsula. 59-65
Muthumary, J., J. A. Jayachandra and M. Bhagavathy Preetha. Development of
conidiomata in the Phyllosticta state of Guignardia mangiferae Roy and observations on
the fine structure of the conidium. 147-155
Nograsek, A. see Scheuer and Nograsek
Olsen, Sigurd, John Haines and Sigmund Sivertsen. Three new hyaloscyphaceous fungi
from Norway and Greenland. 177-182
Pacioni, G. , G. Frizzi, M. Miranda and C. Visca. Genetics of a Tuber aestivum
population (Ascomycotina, Tuberales). 93-100
Parladé, Javier see Alvarez, Parladé, Trappe and Castellano
Phelps, Ryan A. and G. Morgan-Jones. Systematic and biological studies in the
Balansieae and related anamorphs. III. Ascospore and macroconidial germination as
a taxonomic criterion. 41-57
Powell, Martha J. see Blackwell and Powell
Preetha, M. Bhagavathy see Muthumary, Jaychandra and Preetha
Ramaley, Annette W. New fungi from Yucca: Planistromella yuccifoliorum, gen. et sp.
nov., and its anamorph Kellermania yuccifoliorum, sp. nov. and Planistromella
uniseptata, sp. nov., the teleomorph of Kellermania yuccigena. 259-274
Randlane, Tiina and Andres Saag. World list of cetrarioid lichens. 395-403
Reid. Derek A. see Eicker, Van Greuning and Reid
Rogers, Jack D. see Ju, San Martin Gonzdélez and Rogers
see San Martin Gonzalez and Rogers
Ryvarden, Leif and Gast6n Guzman. New and interesting Polypores from Mexico. 1-23
see Hjortstam and Ryvarden
Saag, Andres see Randlane and Saag
Sameva , Ekaterina F. and Ganka G. Bakalova. Septoria thymi sp. nov. from Bulgaria.
387-388
San Martin Gonzalez , Felipe and Jack D. Rogers. Biscogniauxia and Camillea in Mexic«
229-258
see Ju, San Martin Gonzdlez and Rogers
Scheuer, Christian . Lectotypification of Ophiobolus trichellus (Dothideales,
Ascomycetes). 67-69
and A. Nograsek. Trematosphaeria pachycarpa and Herbampulla crassirostris
gen. et spec. nov. (Ascomycetes) 415-424
Sivertsen, Sigmund see Olsen, Haines and Sivertsen
Stierle, Andrea see Strobel, Stierle, Stierle and Hess
Stierle, Don see Strobel, Stierle, Stierle and Hess
Strobel, Gary , Andrea Stierle, Don Stierle and W. W. Hess. Taxomyces andreanae,
a proposed new taxon for a bulbilliferous hyphymycete associated with Pacific Yew
(Taxus brevifolia). 71-80
Sunberg, Walter J. see Banerjee and Sunberg
Toyazaki , Noritsuma see Udagawa, Toyazaki and Tsubouchi
Trappe, James M. see Alvarez, Parladé, Trappe and Castellano
483
Treu, R. Studies on Boletus section luridi. 367-377
Tsubouchi, Haruo see Udagawa, Toyazaki and Tsubouchi
Udagawa, Shun-ichi , Noritsuma Toyazaki and Haruo Tsubouchi. Neosartorya
primulina, anew species of food-borne ascomycetes. 359-366
Upadhyay, H. P. Comments on recent work on Ophiostoma and its synnematous
anamorphs. 411-413
Van Greuning , J. V. see Eicker, Van Greuning and Reid
Venables, Debra A. see Elix and Venables
Visca, C. see Pacioni, Frizzi, Miranda and Visca
Wong, Pak Yau. Type specimens of lichens and lichenicolous fungi in the Canadian
Museum of Nature (CANL). 295-357
Wu, Chi-Guang. Glomales of Taiwan: III. A Compartative study of spore ontogeny in
Sclerocystis (Glomaceae, Glomales). 25-39
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INDEX TO FUNGOUS AND LICHEN TAXA, VOLUME 47
This index includes the names of genera, infrageneric taxa, species, and infraspecific
taxa. New names are in boldface, as are page numbers on which new taxa are
proposed.
Absidia 175
Acanthodochium 219
Acarospora
canadensis 296
punae 296
Acaulospora
scrobiculata 39
Achlya 183, 186, 189, 191
bispora 191
dubia 188-189
Acolium
bolanderi 296
Acrostaphylus 233
Agaricus
brunnescens 97
Ahtiana 395, 403
sphaerosporella 401
wallichiana 402
Albatrellus 23
subrubescens 2, 20
Alectoria
ambigua 296
arctica 297, 312
cornicularioides 297
corymbosa 297
fuscescens 297
glabra 297
imshaugii 297
mexicana 297
pseudofuscens 350
sarmentosa
ssp. sarmentosa 298
stigmata 297
Aleurodiscus 410
aurantius 85
exasperatus 81, 83-85
Allocetraria 395, 403
ambigua 396
cucullata 397, 399
isidiigera 398
nivalis 400
potaninii 400
stracheyi 398, 401
Alternaria 59
Amagdalaria
consentiens
var. japonica 298
continua 298
haidensis 298
Amanita 385, 433, 435-437
subg. Lepidella 436
alauda 436-437
phalloides 433, 435-437
f. phalloides 435
f. umbrina 435
reidii 433-437
verna 437
virosa 437
Amauroderma 8, 21
camerarium 3, 20
Amphinema 407
Amyloathelia
amylacea 83
aspera 81-82
Amylosporus
campbellii 3, 17
Anaeromyces 283
elegans 283-284
mucronatus 283-284
Anaptychia
fragilissima 298
japonica
var. reagens 298
Anisomeridium
nyssaiegenum 299
Anomoporia
albolutescens 3, 17
myceliosa 3
Anthostoma 220, 222
Anthostomella
sinensis 223
Anthracothecium
japonicum 298
thelomorphum 339
Antrodia
gossypina 4, 17
lenis 7
malicola 4, 20
vaillantii 4, 20
Antrodiella
485
486
[Antrodiella] angulatopora 4, 17
versicutis 5, 17
Anzia
afromontana 298
Aphanomyces 183, 192
Arthonia
rufidula 306
Arthopyrenia
submuriformis 298
willeyana 299
Asahinea 395, 402
chrysantha 397, 399, 401
scholanderi 399, 401
Ascochyta 148-150
cypericola 76
Aspergillus 365
sect. Fumigati 365
Aspicilia 318
quartzitica 299
Asteromella
brassicae 132
Atkinsonella 41, 56-57
hypoxylon 43, 54, 56
var. texensis 56
texensis 43, 56-57
Aurificaria
liteo-umbrina 5, 20
Bacidia
colchica 299
finkii 300
lisowskii 299
nivalis 299
scutellifera 299
spirospora
var. patagonica 299
vasakii 299
violascens 300
Baeomyces
weberi 300
Bahianora
poeltii 300
Bainieria 412
Balansia 41-43, 56-57
aristidae 41, 43-44, 46, 48-50, 56-57
atramentosa 56
claviceps 42, 57
epichloe 41, 43-44, 50, 52-54, 57
henningsiana 57
linearis 42
obtecta 42
strangulans 41, 57
Balansiopsis 56
Balsamia 214
vulgaris 203
Basidiobotrys 233, 241
Belemnospora 92
Bilimbia
finkii 300
Biscogniauxia 226-227, 229-231, 233-
234, 238
atropunctata 231-232, 240
cinereo-lilacina 236
comedens 231
divergens
var. macrospora 231, 233
exutans 238
fuscella 229, 231, 233
grenadensis 231, 236
mediterranea 232, 236
nummularia 231-232, 238-240
var. exutans 232, 238
var. merrillii 232, 238-239
var. nummularia 239
var. "pseudopachyloma" 232, 239-
240
sp. 231-232, 234, 236, 240
Blumeria
graminis 197
Boletus 367-368, 375-376
sect. Luridi 367, 376-377, 385
calopus 376-377
piedmontensis 376
pseudo-olivaceus 367, 369-371, 374,
376
queletii 376
roseobadius 367, 371-374, 376
rufocinnamomeus 367, 372-373, 376
subluridellus 367, 369, 373-377
subvelutipes 367
Botanamphora 415, 419
pachycarpa 415-416, 419
Botryodiplodia
theobromae 150
Botryosphaeria
dothidea 271
Brevilegnia 184, 186, 188, 191-192
bispora 186, 191
unisperma 188
var. litoralis 188
Bryoria
carlottae 300
cornicularioides 297
friabilis 300
furcellata 398
fuscescens 297
glabra 297
[Bryoria] pikei 300
pseudocapillaris 301
salazinica 301
spiralifera 301
trichodes
subsp. americana 297
subsp. trichodes 343
Buellia
excellens 301, 340
galapagona 301
imshaugii 301
multispora 301
rivas-martinezii 301
tephrodes 301
Buelliella
eximia 302
physciicola 302
Buergenerula 271
Bulbothrix
microscopica 101-102, 104
queenslandica 101, 126
subtabacina 101-102, 127
Burgoa 76
Byssocorticium 407
coprophilum 407
lutescens 407
Byssoloma
amazonicum 302
anomalum 302
Caecomyces
equi 293
Cainia
graminis 422
Calicium
adspersum
ssp. australe 302
constrictum 302
cryptocroceum 302
curtisii
var. splendidula 302
fuscipes 303
obscurum 303
parvum 303
Caloplaca
bisagnonis 303
brattiae 303
britannica 303
citrina
var. arcis 303
coralloides 334
fraxinea 303
hensseniana 303
487
litoricola 304
marina
ssp. americana 304
sbarbaronis 304
sorediata
var. tenuis 304
spotornonis 304
trabicola 304
Calyptralegnia 186
achlyoides 187
Camillea 227-229, 233, 241, 247, 254-
phe:
broomeiana 250
cyclisca 243
flosculosa 250
fossulata 236, 242-243, 250
gigaspora 242-243, 248
guzmanii 229, 243-244, 246, 248
hainesii 242, 246
hyalospora 246
labellum 242, 244, 246-247
magnifica 229, 234, 243, 246-247,
248
mexicana 229, 234, 242, 248, 250-
251
mucronata 242, 250-251
obularia 242, 251
punctulata 242, 250-251
stellata 242, 252
tinctor 243, 252
verruculospora 241, 253
Candelariella
canadensis 304
faginea 304
hudsonica 304
lambii 304
plumbea 305
Canomaculina
melanochaeta 101, 127
Canoparmelia 103, 105
herveyensis 101-102, 103-104
macrospora 103
owariensis 101, 127
pruinata 103
pustulescens 101, 127
raunkiaeri 102-103
subarida 101, 103-104
whinrayi 101, 104-105
Catapyrenium
caeruleopulvinum 305
tuckermanii 312
Catillaria
corymbosum 297
italica 305
488
[Catillaria] melanopotamica 305 delavayi 397
Ceratocystiopsis 411, 413 delisei 397-398
Ceratocystis 411-413 denticulata 397
davidsonii 412 dermatoidea 397
piceae 412 diffusa 397
sagmatospora 411 elenkinii 397
ulmi 412 endocrocea 398
Cetraria 395, 403 endoxantha 398
aculeata 396 endoxanthoides 398
agnata 396 epiphorella 398
alaskana 305, 396 ericetorum 397-399, 401
albopunctata 396 eriophylla 398
aleurites 396 everniella 398
alvarensis 396 fahlunensis 398
ambigua 396 fallax 398
americana 396 fastigiata 398
andrejevii 396, 401 fendleri 398
annae 396 f. coralligera 305
antarctica 396 formosana 398
arborialis 396 furcellata 398
arctica 396 gilva 398
arenaria 396 glauca 398
asahinae 396 globulans 398
atlantica 396 gracilenta 398
aurescens 396 halei 398
australiensis 305, 396 hepatizon 398, 400
bavarica 396 herrei 398
billardieri 396 hiascens 398
bohemica 396 hypotrachyna 398
braunsiana 396 iberica 305, 398
californica 396-397 idahoensis 398
canadensis 397 inermis 398
caperata 397 inflata 398
capitata 397 isidioidea 399
chicitae 397 islandica 399-400
chlorophylla 397 var. arborialis 305
chrysantha 397 japonica 399
ciliaris 397 juniperina 399
citrina 397 kamczatica 399
clarkii 397 komarovii 399
f. clarkii 397 kurodakensis 399
collata 397 kurokawae 399
commicta lacera 399
f. sorediella 401 lacunosa 399
commixta 397-398 laevigata 399
complicata 397 lanata 399
coralligera 305, 397 laureri 399
corallophora 397 laxa 399
corniculata 397 leucostigma 399, 401
corrugis 397 libertina 399
crispa 397 madreporiformis 399
cucullata 397 magnussonii 399
culbersonii 397-398 megaleia 399
daibuensis 397 melaloma 399-400
[Cetraria] merrillii 399
microphylla 399
microphyllica 399
muricata 399
nepalensis 399
nephromoides 399
nigricans 397, 399
nigricascens 396-397, 399-401
nipharga 399
nipponensis 399
nivalis 400
normoerica 400
norvegica 400
nova-zelandiae 400
nuda 400
oakesiana 400
ochrocarpa 400
odontella 400
orbata 400
ornata 400
pachysperma 400
pallescens 400
pallida 400
pallidula 400
perstraminea 400
pinastri 400
placorodia 400
platyna 400
platyphylla 400
platyphylloides 400
polyschiza 400
potaninii 400
pseudocomplicata 400
pubescens 400
ramulosa 400
rassadinae 400
regenerans 400
rhizophora 400
rhytidocarpa 401
richardsonii 401
rugosa 401
sanguinea 401
saviczii 401
scholanderi 401
scutata 401
sepincola 401
septentrionalis 401
sibirica 401
sikkimensis 401
simmonsii 401
sorediella 401
stenophylla 401
stracheyi 401
straminea 401
489
stuppea 401
stygia 401
subalpina 306, 396, 401
subfendleri 401
subscutata 401
sulphurea 401
tenuifolia 401
tenuissima 401
terrestris 401
teysmanni 401
thomsonii 402
thyreophora 402
tilesii 402
togashii 402
tristis 402
tubulosa 402
tuckermanii 402
ulophylla 402
ulophylloides 402
urceolata 402
viridis 402
wallichiana 402
weberi 402
xizangensis 402
yunnanensis 402
Cetrariopsis 395, 403
wallichiana 402
Cetrelia 395, 403
alaskana 305, 396
braunsiana 396
cetrarioides 397
chicitae 397
collata 397
davidiana 397
delavayana 397
isidiata 398
japonica 399
monachorum 399
nuda 400
olivetorum 396, 400-401
orientalis 400
pseudocollata 400
pseudolivetorum 400
sanguinea 399, 401
sinensis 401
Cetreliopsis 395
rhytidocarpa 399, 401
Chaenotheca
brunneola 303
Chaenothecopsis
nana 306
nigropedata 306
sagenidii 306
tasmanica 306
490
Chaetocalathus 379, 384-385
sect. Holocystis 384
liliputianus 384
magnus 379, 382-384
Chaetoconis 272
"Chaetonaevia
archangelica" 180
Chaetospermella 62
Chaetospermum 59, 62-64
camelliae 63-64
carneum 63
chaetospermum 64
chaetosporum 59-60, 62-64
gelatinosum 62-63
tubercularioides 63
Charcotia
rufidula 306
Cheiromycina
petri 306
Chithramia 64
Choiromyces
magnusii 204
meandriformis 204
venosus 204
Ciliochorella
mangiferae 150
Ciliospora 62
Cladidium
bolanderi 318
Cladina
arbuscula
subsp. beringiana 307
boliviana 307
conspicua 307
terrae-novae 309
f. cinerascens 309
stygia 309
Cladonia
alpestris
f. aberrans 307
anitae 307
arbuscula
subsp. beringiana 307
boliviana 307
botrytes 307
convoluta
var. vagans 307
crispata
var. infundibulifera
f. albopunctata 307
var. cetrariiformis 307
favillicola 307
gracilis
subsp. vulnerata 307
hedbergii 308
kauaiensis 308
labradorica 308
libifera 308
magyarica 308
norvegica 308
patagonica 308
physodalica 308
poeciloclada 309
pollicum 308
rangiferina
f. caerulescens 309
f. leucosticta 309
squamosa 308
subchordalis 309
terrae-novae 309
f. cinerascens 309
thomsonii 309
wainii 309
Cladorrhinum 74, 76
foecundissimum 74
Cladosporium 59, 146
herbarum 142
Clinostomum
luteolum 309
vitellinum 309
Coccocarpia
stellata 309
Coccotrema
maritimum 310
Coelocaulon
aculeatum 396-397
muricatum 399, 401
Coelopogon
epiphorellum 398
Coleophoma
cylindrospora 150
Collema
leptaleum 310
mircoptychium 310
Collybia 385
subg. Rhodocollybia 385
Coltricia 13
Combea
californica 342
Confertobasidium 410
Coniella
fragariae 150
Coniocybe
amabilis 310
Coniothecium 431
Conotremopsis
weberiana 310
Coriolopsis
[Coriolopsis] caperata 7
caperatiformis 7
floccosa 6, 20
rigida 6, 20
Coriolus
ochrotinctellus 5
Cornicularia 403
normoerica 400, 402
Coronicium 410
Corticium
amylaceum 83
coprophilum 407
filium 409
vallum 83
Cortinarius 461, 465
subg. Leprocybe 464-465
sect. Limonei 464-465
sect. Orellani 465
lewisii 462, 464-466
limonius 464-465
nothosaniosus 464-465
Creosphaeria 219, 222
pinea 222
riograndensis 222-223
sassafras 219, 222-223, 224, 226-227
verruculosa 222
Crinipellis 379, 382, 385
sect. Iopodinae 382
sect. Ionopodinae 382
hygrocybioides 382
insignis 382
purpurea 382
ticoi 379, 380-382
Cristinia 405-407, 409-410
artheniensis 405-406
brevicellularis 405-406
coprophila 405-407
filia 409
gallica 406-407
helvetica 405-406
mucida 407
rhenana 405-407, 408
sasae 405
sonorae 409
Crocicreas 178
Cryptophiale 87, 91-92
aristata 91
caudata 87-88, 91
enormis 91
kakombensis 91
novae-caledoniae 87, 89-91
secunda 91
Cryptophialoidea 91-92
Cytospora
491
turgida 220
Dacryobasidium 405, 407
Dactylina
arctica 396
beringica 310
madreporiformis 399
ramulosa 400
Daedalea
ambigua 8
elegans 6
microsticta 6, 19-20
Daldinia 254
Datronia
caperata 7, 20
Delastria 214
rosea 204
Dentipellis
fragilis 407
Dermatocarpon
corticola 310
"corticolum" 310
gorzegnoense 310
vagans 310
Diatrype
hypophloea 226
stigma 224
Dictyuchus 186
clavatus 184-185, 187
Didymella 271
Dilophospora
alopecuri 64
Dimerella
chiodectonoides 310
degeneri 311
frederici 311
isidiigera 311
pocsii 311
subdentata 311
subfallaciosa 311
usambarensis 311
Dingleya 217
Diplomitoporus
lenis 7, 17
Diploschistes 129
actinostomus 312
conceptionis 312
hypoleucus 312
montevidensis 312
sbarbaronis 312
straminescens 312
Dirina
californica 312
492
Dolichocarpus
chilensis 312
Drechslera 59
Elaphomyces 201
anthracinus 204
granulatus 204
muricatus 201, 204
personii 204
trappei 204
Endocarpon
mailae 312
tuckermanii 312
Endogone
microcarpa 205
Entomopatella 62
Entophylyctis
variabilis 289, 292
Eopyrenula
intermedia 338
Ephelis 41-44, 48, 50, 52, 56
mexicana 42
Epichloe 41
typhina 43, 54, 56-57
Erioderma
knightii 398
Erysiphe
convolvuli 194, 197
Esslingeriana 395
idahoensis 398
Eutypa
armeniaceae 228
Evernia
perfragilis 297, 312
stracheyi 401
Everniastrum
fragile 312
Fellhanera
endopurpurea 312
Flavodon
flavus 7, 17
Flavoparmelia 129
euplecta 105-106
kantvilasii 101, 105-106, 109
Fomes
ohiensis 11
Fomitopsis
cubensis 11
melliae 8, 17, 19
Fulgensia
canariensis 313
Gaertneriomyces 292
Ganoderma 23
coffeatum 8, 19-20
Gautieria
mexicana 204
morchelliformis 204, 214
Genea 214
klotzschii 204
verrucosa 205, 214
Geniculosporium 219
Geolegnia 186
Geopora
arenicola 205
cervina 205
cooperi 205
foliacea 205
nicaensis 205
sumneriana 205
tenuis 205, 215
Glomus 25-26, 34, 201
ambisporum 26, 34, 39
constrictus 39
dimorphicum 34
heterosporum 25, 34, 39
macrocarpum 201, 205
microcarpum 205
Glyphis
achariana 313
Gonohymenia
cribellifera
ssp. macrocarpa 313
lusitanica 313
undulata 313
Grammothele
fuligo 8, 20
Graphilbum 411-412
Graphina
aibonitensis 313
muscicola 313
subvirginalis 314
Graphis
subvirginalis 313
Graphium 411-412
penicilloides 412
Guignardia
mangiferae 147, 149
Gyalectidium
colchicum 314
Gyalideopsis
alnicola 314
kalbii 314
trapperi 314
[Gyalideopsis] vezdae 314
Gyrophoropsis
dwaliensis 314
Haematomma
inexpectatum 314
pustulatum 314
Hafellia
fosteri 315
Heppia
uruguayensis 315
Herbampulla 419, 422
crassirostris 415, 419, 421-422, 424
Heterodermia
desertorum 315
stichensis 315
Heteropatella 272
Heterothecium
pachycheilum 315
Homothecium
sorediosum 315
Hubbsia
californica 315
lumbricoides 315
Huilia
circumnigrata 315
Humphreya 8
Hyalopesotum 411
Hyalopeziza 180
subg. Unguicularia 180
subg. Unguiculella 181
citrinus 206
muticus 201, 206
niveus 201, 207
olivaceus 207
populetorum 207
remyi 207
sp. No. 1 206
sp. No. 2 206
tener 201, 207
vulgaris 207
Hyperphyscia
mobergii 316
Hyphoderma 410
Hyphodontia
subs. Ellipsosporae 409
Hypocenomyce
foveata 275, 277, 279
leucococca 316
Hypochniciellum 407
Hypogymnia
austerodes 330
billardieri 281, 396
heterophylla 316
hokkaidensis 316
kiboensis 316
mollis 316
oceanica 316
rugosa 330
Hypotrachyna 275, 280
booralensis 101, 106, 109
densirhizinata 106-107
heterochroa 107
archangelica 177-178, 180 immaculata 275, 277-279
groenlandica 177-178, 181 livida 275, 277
raripila 180 osseoalba 275, 279
Hydnangium 158 proserpinensis 101, 107-109
aurantiacum 205 pustulifera 108
carneum 206 subpustulifera 101, 108-109
carotaecolor 205
pila 157-158, 206
Hydnocystis
clausa 214
Hydnotrya
tulasnei 206
Hydnum
helveticum 405, 407
mucidum 407
Hydrothyria
venosa 316
Hymenogaster 201
albus 201, 206
bulliardi 206
calosporus 206
cerebellum 206
virginica 331
Hypoxylon 219, 222, 226, 228, 230,
234, 238-241, 254-255
sect. Entoleuca 228
atropurpureum 234
bifrons 223
callostroma 223
cinereo-lilacinum 234, 236
cinereum 225-226
comedens 233
diathrauston 222
divergens 233
var. macrospora 233
flosculosum 250
fossulatum 250
fuscellum 233
494
{[Hypoxylon] glycyrrhiza 246, 254
grisea 226
hainesii 246
hemisphaericum 244, 248
hypophlaeum 224, 226
maculum 239
microplacum 224, 226-227, 232, 238
nummularium 231, 239
var. exutans 238
var. merrillii 238
var. pseudopachyloma 239
var. theissenii 241
punctulatum 251, 254
sassafras 222-223
tinctor 233
valsarioides 223
Hysterangium 201, 215
clathroides 201, 207
gardneri 201, 207
membranaceum 210
rickenii 207
separabile 207
stoloniferum
var. rubescens 207
thwaitesii 207
Immersaria
carbonoidea 322
Imshaugia
evernica 101, 127
placorodia 397, 400
Incrustoporia
nivea 9, 20
Inocybe
geophylla 392
Inonotus
hispidus 9, 20
munzii 9, 20
Involucrothele
ligurica 316
Ionaspis
ventosa 317
Jenmania
osorioi 317
Jongiella 254
Juleila
fallaciosa 335
Junghuhnia
nitida 10, 20
Kalbiana
brasiliensis 317
Karsteniomyces 425-426, 428
llimonae 425-426, 427, 429, 431
peltigerae 426
tuberculosus 426
Kellermania 65, 259, 265-266, 268,
272-274
anomala 265-266, 270, 273
yuccifoliorum 259-260, 262, 265-
266, 268-273
yuccigena 259, 265, 267-268, 270,
272-273
Kiessleriella 271
Kochiomyces 292
Labyrinthomyces 217
donkii 208, 215
"Laetinaevia
archangelica" 181
Laetiporus
sulphureus 14-15
Lecanactis
chloroconia 317
premnea
var. chloroconia 317
subdryophila 317
Lecania
brialmontii 349
constricta 317
macrocarpa 317
sbarbaronis 317
Lecanora
aeruginascens 317
austrooceanica 318
bolanderi 318
caesiorubella
subsp. merrillii 318
caesiosora 318, 321
californica 318
christoi 318
cinerofusca
var. appalachensis 318
concinna 318
"concinnum" 318
congesta 318
freyi 319
fuliginosa 319
geophila 321
gisleri 319
homalea 319
imshaugii 319
insolata 319
{[Lecanora] kutakii 319
laatokkaensis 343
luteovernalis 319
mellea 320
microbola 320
"olea" 320
oleae 320
opiniconensis 320
orae-frigidae 320, 322
pachysoma 320
perplexa 320
pseudopinguis 320
pycnocarpa 321
salicicola 321
soralifera 318, 321
superfluens 321
texana 321
thallophila 321
utahensis 321
weberi 321
Lecidea
(Psora) 322
baffiniana 321
carbonoidea 322
grassiana 322
gypsicola 322
hyotheja 322
leproloma 322
patagonica 322
santensis 322
shushanii 322
sorediata 320, 322
ullrichii 323
Lempholemma
dispansum 323
Leprocaulon
arbuscula 343
pseudoarbuscula 347-348
subabicans 349
Leproloma
diffusum
var. chrysodetoides 323
Leptogium
byssinum 323
caesiellum 323
Leptographium 412
Leptoguignardia 271
Leptosphaeria 178, 271, 416
hazslinskyana 415-416
pachycarpa 415-417, 424
Letharia
togashii 323
wandelensis 323
Leucogaster
floccosus 208, 215
Libertella 222
Lichen
candelarius 323
cavernosus 397
salazinus 323
Lichenoconium
parasiticum 323
Lichenostigma
maureri 324
Lichina
macrospora 324
polycarpa 324
rosulans 324
Lindquistia 219
Lobaria
fulva 324
fuscotomentosa 324
gyrophorica 324
oregana 350
tuberculata 324
Loculotuber 201
gennadi 201, 208, 214
Lopadostoma 219
495
turgidum 219-220, 222, 226-227
var. minus 220
Loweporus 11
Macrophomina
phaseolina 148-150
Marasmius
hygrocybioides 382
Maronella
laricina 324
Martellia
mediterranea 157, 208
mistiformis 157
pila 157
Masonhalea 395
richardsonii 401
Mastigonema 62
Mazosia
pilosa 325
pseudobambusae 325
Megalospora
foersteri 325
pachycheila 315
Melanaria
macounii 325
var. meizotoca 325
Melanelia 403
albertana 328
stygia 401
496
[Melanelia] tominii 330
trabeculata 331
Melanogaster
ambiguus 208
broomeianus 208
variegatus 208
Melaspilea
fuscolimitata 325
Melogramma 422
Micarea
muhrii 325
vulpinaris 325
Microdochium
fusarioides 449
Microporellus
obovatus 10, 20
Microsphaeria
euonymi-japonici 195
trifolii 195
Microthelia
atramentea 325
Milospium
graphideorum 430
Minimedusa 76
Monochaetiella
hyparrheniae 272
Monochaetiellopsis 272, 274
Mucor 176
Muhria
urceolata 325
Mycocalicium
fuscipes 303
Mycoglaena
quercicola 325
Mycosphaerella 131-133, 135, 142,
144-146
allicina 142
alyssi 132, 144-145
brassicicola 131-132, 144-146
carniolica 133, 144-145
confinis 133-134, 144-145
cruciferarum 131, 134, 137, 144-145
denigrans 131, 135-136, 144-145
densa 131, 136, 144-145
hambergii 136, 144-145
isatidis 137, 144-145
napicola 144-145
pachyasca 131, 137-139, 144-145
pashkiensis 131, 139-140, 144-145
phlomidicola 135
pyrenaica 131, 136, 141, 144-145
tassiana 131, 133-134, 136, 138-140,
142-145
vesicariae-arcticae 142-145
Myriogenospora 56
atramentosa 41-44, 46, 54, 56-57
Nectria 43, 47
aquifolii 43
coryli 43
Neocallimastix 292
frontalis 289, 292-293
variabilis 293
Neofuscelia 112, 115
delisea 112
imitatricoides 101, 110-111, 113
imitatrix 110-111
incantata 110
kondininensis 101, 110, 113
loxodella 111-112
luteonotata 112, 114
melanobarbatica 115
parasitica 101, 111-113
parviloba 114-115
pulla 111
pyrenaica 112
scabrella 114
scabrosina 101, 112-114
subbarbatica 101, 114, 117
subincerta 112, 128
verisidiosa 111-112
Neosartorya 359-360, 364-365
fischeri 364-365
hiratsukae 359, 364, 366
primulina 359-360, 361-362, 364
pseudofischeri 365
Nephroma
americana 396
chubutense 326
kuenhnemannii 326
lepidophyllum
f. "hypomelaena" 326
f. hypomelaenum 326
occultum 326
Nephromopsis 395, 403
asahinae 396
californica 397
delavayi 397
ectocarpisma 397, 399
endocrocea 397-398
endoxanthoides 398
globulans 398
isidioidea 399
komarovii 399-400
kurokawae 399
laureri 397, 399
laxa 397, 399
[Nephromopsis] morrisonicola 399
nipponensis 399
ornata 398, 400
pallescens 397, 400-401
pseudocomplicata 400
rugosa 401
stracheyi 401
yunnanensis 402
Neuropogon
rohmederi 326
f. ushuaiensis 326
trachycarpus
f. elatior 326
Nodulisporium 219, 229-230, 233-234,
252
Nozemia
syringae 439
Nummularia 230, 241, 254
broomeiana 254
fuscella 233-234
"Nummularia"
grisea 226
hypophloea 226
sp. 219, 224, 226-227
viridis 219, 224, 226
Numulariola 233, 238, 254
atropunctata 232
broomeiana 251
exutans 238
mediterranea 236
pseudopachyloma 240
Ochrolechia
gowardii 326
juvenalis 326
montana 327
pseudopallescens 327
subisidiata 327
trochophora
var. pruinirosella 327
tuckermanii 327
yasudae 327
Octaviania
asterosperma 208
pila 206
Oidium 193-194, 196-197, 199
Oligoporus
fragilis 10, 20
Omphalodina
bullata 327
Opegrapha
albidoatra 327
astraea 327
497
pertusariicola 327
subabnormis 328
Ophiobolus
trichellus 67-68
Ophiochaeta
trichella 68
Ophiostoma 411, 413
davidsonii 412
sparsum 412
Oropogon
colibor 328
diffractaicus 328
mexicanus 328
Orpinomyces
bovis 292
Ovularia
syringae 439
Pachnodium 411
Paecilomyces 365
Pannaria
leucostictoides 328
Papulaspora 76
Paraparmelia 129
alabamensis 328
lithophiloides 115
yamblaensis 101, 115, 117
Paraphaeosphaeria 271
Parmelaria 395
subthomsonii 401
thomsonii 402
Parmelia
subg. Xanthoparmelia 128
alabamensis 328
albertana 328
ambigua
var. halei 328
cetrarioides 397
chlorochroa 328
crinita
f. varians 328
crowii 101, 116-117
cryptoxanthoides 329
dubia
var. scrobiculata 329
elabens 329
erumpens 116
formosana 281
frondifera 329
frostii 398
gloriosa 329
haitiensis 127
hygrophila 329
498
[Parmelia] jejunga 118
livida 281
melanochaeta 127
monachorum 399
multipartita 126
olivetorum 400
omphalodes
subsp. glacialis 329
owariensis 127
pacifica 329
physodes
var. enteromorpha
f. rugosa 329
f. subisidioides 330
pseudolivetorum 400
pseudotenuirima 116
pseudovirens 127
pustulescens 127
queenslandica 126
ralla 330
reptans 330
rubsecens 401
saximontana 330
semansiana 330
skultii 329
sphaerosporella 401
spinibarbis 330
squarrosa 330
subdistorta 330
subtabacina 127
sulcifera 331
trabeculata 331
virginica 331
xanthosorediata 127
Parmeliella
concinna 331
granulata 331
Parmelina 118, 129, 425
conlabrosa 118-119
endoleuca 119
euplectina 101, 116-118
johnstoniae 101, 118-119, 121
labrosa 105
pastillifera 425, 428-429
pseudorelicina 116, 118
quercina 425-426, 429
tiliacea 425, 428, 430
Parmelinopsis
evernica 127
minarum 119
protocetrarica 101, 119, 121
subambigua 328
Parmentaria
astroidea 338
Parmotrema 402
demethylmicrophyllinicum 331
indicum 331
merrillii 120
perforatum 397
pseudovirens 101, 127
queenslandense 331
submerrillii 101, 120-121
subtinctorium 328
zollingeri 120
Peltigera 426
kristinssonii 331
malacea
f. hasimotoi 332
pacifica 332
Peltula
cylindrica 332
Penicillium 365
Perenniporia
ohiensis 11, 20
tephropora 11, 17
Pertusaria
glaucomela 333
gymnospora 332
macounii 325
neoscotica 332
panyrga 333
paramerae 332
santamonicae 332
suboculata 332
subpupillaris 332
subvelata 333
sulcata 333
victoriana 333
Pesotum 411-412
ulmi 411
Pestalotiopsis
palmarum 150
Pestalozziella 65
Phaeocalicium
asciiforme 333
curtisii 303
Phaeocollybia 385
Phaeolopsis
veraecrucis 14
Phaeopeccania
australiensis 333
hispanica 333
Phaeophyscia
cernohorskyi 334
spinellosa 333
Phaeosphaeria 424
Phellinus 12
chryseus 11, 17
[Phellinus] contiguus 12, 17
ferrugineo-velutinus 12, 17
gilvus 19
grenadensis 12, 17
rhytiphloeus 12, 17
rimosus 13, 20
Phialocephala 412
Phialographium 411-412
sagmatosporae 411
Phloeophthora
syringae 439
Phlyctis
pulveracea 333
Phlyctochytrium
arcticum 292
Phomatospora 422
Phylloporia
chrysita 13, 20
frutica 13
spathulata 2, 13-14
veraecrucis 2, 14, 20
Phylloporus 385
boletinoides 385
Phyllopsora
corallina 322
Phyllosticta 147, 149-150, 154
Physcia
duplicorticata 333
hirsuta
var. echinella 334
Phytophthora 163, 175, 440-441, 447-
452
cactorum 440, 448, 452
cinnamomi 446
citrophthora 449-450
crytogea 449
hibernalis 439-451, 453-457, 460
megasperma 445, 449
parasitica 446
porri 445, 450
syringae 439-457, 460
Picoa
juniperi 208
Pilidiella
quercicola 150
Pilophorus
cereolus
var. hallii 334
clavatus 334
Piptarthron 259, 265-266, 272-273
pluriloculare 274
uniloculare 259, 265, 274
Piptoporus
soloniensis 14, 17
499
Piromonas
communis 285-286, 293
minima 286
Piromyces 285-286, 289, 292
communis 286, 292
dumbonica 286, 293
mae 292-293
minutus 285-286, 287-288, 290, 292
rhizinflata 286, 293
Placodium
coralloides 334
Placopsis
alphoplacoides 334
var. clavidera 334
contortuplicata
f. fuegiensis 334
dusenii 334 ;
gelidoides 334
lateritioides 334
perrugosa
f. activa 335
salazina 335
trachyderma 334
var. Clavifera 334
Placynthium
petersii 337
Plagiocarpa
septemseptata 335
Planistroma 259, 271, 273
obtusilunatum 265, 273-274
yuccigena 274
Planistromella 259-260, 271, 273
uniseptata 259, 264, 267-268, 271
yuccifoliorum 259-260, 261-262,
264-267, 271
Platismatia 395, 403
erosa 398, 400
formosana 398
glauca 398
herrei 398, 402
interrupta 398
lacunosa 397, 399
norvegica 399-400
regenerans 400
stenophylla 401
tuckermanii 396, 402
Platysma
thomsonii 402
subperlatum 401
Piuteus 389-390, 394
sect. Celluloderma 389
sect. Hispidoderma 389
sect. Pluteus 389-390, 392
cervinus 390
500
[Pluteus] cinerascens 389-390 Psilolechia
heterocystis 389-390, 392 leprosa 336
laricinus 389-390, 392 Psora
magnus 390 cerebriformis 336
salicinus 393-394 montana 337
var. achloes 394 pacifica 337
var. americanus 389-390, 393-394 texana 337
var. salicinus 394 Psoroma
Polyblastia internectens 337
cucurbitula 335 Psorotichia
sbarbaronis 335 rimosa 336
Polyblastiopsis Pterygiopsis
quercicola 335 australiensis 337
Polycauliona canariensis 337
coralligera 335 convoluta 337
Polyporus 13 Pterygium
caperatus 7 petersii 337
elegans 15 Punctelia
pargamenus 16 bolliana 329
pseudosulphureus 15 perreticulata 329
semipileatus 9 rudecta 397
soloniensis 15 semansiana 330
varius 15, 20 Pustularia
veraecrucis 14 insignis 215
Polystictus 13 Pyrenastrum
Poria 22 gemmeum 337
myceliosa 3 Pyrenopsis
vaillantii 4 sojakii 338
vincta 15 Pyrenula
Porina cinerella
corruscans 335 var. quadriloculata 338
mangiferae 335 leucoplaca
Porocyphus
kalbarrensis 335
Porogramme
fuligo 8
Porpidia
carlottiana 335
thomsonii 336
Protoparmelia
laricata 336
Pseudephebe
pubescens 399-400
Pseudocyphellaria
anomala 336
coriacea 341
exanthematica 336
hirsuta
f. leucosticta 336
rainierensis 336
Pseudoidium 194
Pseudoparmelia
aradensis 396
crystallicola 336
var. pluriloculata 338
macounii 338
maculata 338
neglecta 338
subsp. occidentalis 338
nitidella
var. maculata 338
occidentalis 338
pseudobufonia 338
santensis 355
shirabeicola 338
thelomorpha 338
Pyroporellus
subblinteus 5
Pythium 163, 175, 450
Pyxine
daedalea 339
Radulodon 409
americanus 409
Radulum
[Radulum] gallicum 407
mucidum 407
Ramalina
americana 339
atlantica 339
bicolor 339
cactacearum 339
euxini 339
inflata 398
sideriza 339
superfraxinea 339
tenuis 339
tigrina 340
Ramonia
intermedia 340
kandlerii 340
Reddellomyces 217
Relicina 128
Rhizocarpon
alaxensis 340
atlanticum 340
barilochense 340
compositum 301, 340
constrictum 340
cumulatum 340
hensseniae 340
inimicum 340
renneri 341
Rhizophlyctis
rosea 289, 292
Rhizoplaca
bullata 327
Rhizopogon 201, 212
aestivus 209
colossus 215
luteolus 208
provincialis 209
roseolus 209, 213
rubescens 209
subareolatus 201, 209, 213
ventricisporus 201, 209
vinicolor 215
vulgaris 209
Rhizopus 161-163, 175-176
arrhizus 175
azygosporus 163
homothallicus 162
microsporus 162, 175-176
oryzae 163, 175-176
sexualis 162
stolonifer 162, 176
Rhodocybe 384
Ricasolia
beckettii 341
Rigidoporus
vinctus
var. cinerea 15, 17
var. vinctus 15
Rimeliella
haitiensis 101, 127
Rimularia
fuscosora 341
Rinodina
afghanica 341
ascociscana 341
calculiformis 341
conradii 342
dakotensis 341
humilis 341
milliaria
f. obscura 341
oregana 342
pallida 342
sabulosa 342
Robillarda
depazeoides 150
Rocella
galapagoensis 342
Rocellina
luteola 342
olivacea 342
Rosellinia
diathrausta 222
linderae 223
prinicola 223
Rozites 385
Ruminomyces 283
elegans 283-284
Sagenidiopsis
merrotsii 342
Saprolegnia 183, 192
Sarrameana
tasmanica 342
Sawadaea 197
Schismatomma
californicum 312
Schistophoron
variabile 342
Schizopelte
californica 342
Sclerococcum 425, 430-431
epiphytorum 430
serusiauxii 425, 428-430
sphaerale 430
Sclerocystis 25-26, 28, 30, 32, 34, 36,
38-39
501
502
[Sclerocystis] clavispora 25-28, 30, 32, Stachybotrys 92
36, 39 Stagonospora 271-272
coremioides 25-28, 30, 32, 34, 36, 38 gigantea 259, 271
liquidambaris 25-28, 30, 34, 36 Stephanocyclos
rubiformis 25-27, 30, 32, 34, 36, 38 henssenianus 343
sinuosa 25-28, 30, 32, 34, 36, 38 Stereocaulon
taiwanensis 25-28, 30, 32, 34, 36, 38 alpinum
Scoleconectria var. erectum 343
cucurbitula 43 arbuscula
Scolecosporiella 272 var. aberrans 343
Scutula 426 argodes 343
aggregata 426 argus 343
miliaris 426 var. stenospermum 344
Septoria 150, 387
thymi 387-388
azulense 344
botryosum
Sepultaria f. depressum 344
arenicola 205 f. pygmaeum 344
arenosa 205 capense 344
foliacea 205 colensoi
Setaria var. raegens 344
trichodes 343 condensatum
Skeletocutis var. sorediatum 344
nivea 9 coralloides 344
Skyttea var. flabellatum 344
tephromelarum 343 var. occidentale 345
Sphaerella cornutum
alyssi 132 var. corallizans 345
brassicicola 132 corticatulum
carniolica 133
confinis 133-134, 136
var. hambergii 136
cruciferarum 134
denigrans 135
densa 136
napicola 137
pachyasca 137, 139
pyrenaica 141
tassiana 138, 142
vesicariae-arcticae 142
Sphaeria
var. capense 344
var. complanatum 345
var. procerum 345-346
crambidiocephalum 345
dactylophyllum 344
var. flabellatum 344
var. occidentale 345
dendroides 345
denudatum
var. umricola 345
var. "umbricolum" 345
depressum 344, 346
brassicicola 132 dusenii 345
callostroma 223-224 esterhuysenae 345
cruciferarum 134 evolutoides
prinicola 223 var. paschaleoides 345
sassafras 222-224 exutum 346
turgida 220 fastigiatum
Sphaerophorus var. dissolutum 346
notatus 343
ramulifer 343
Spizellomyces 289, 292
f. confluens 346
glabrum 344, 348
f. flabellans 347
Sporothrix 411-412 glareosum
Squamaria var. brachyphylloides 346
configurata halei 346
f. diffracta 343
intermedium
[Stereocaulon intermedium] f.
compactum 346
japonicum
var. subfastigiatum 346
var. tokioense 346
leprocauloides 346
melanopotamicum 345-346
meyeri
f. tucumanum 346
montagneanum 347
myriocarpum
var. altaicum 347
nanodes
f. schadeanum 347
novoarbuscula 347
octomerelloides 347
octomerellum 347
papuanum 347
paschale 348
subsp. evolutoides
f. laxatum 347
f. paschale 347
var. alpinum
f. flabellans 347
var. serpens 348
v. evolutoides
f. sorediatum 348
patagonicum 348
f. subirregulare 348
philippinense 348
pileatum 348
var. nipponicum 348
f. macrum 348
proximum
var. gracilius 348
pseudoarbuscula 348
pseudomassartianum 349
pygmaeum 349
ramulosum 350
var. gracilius 348
var. pulvinare
f. crebratum 349
salazinum 323
saxatile
f. paschaleoides 346
f. sorediatum 348
spathuliferum
f. congestum 346
f. pygmaeum 344
speciosum 349
var. surreptans 349
stenospermum 344
subcoralloides
f. sorediascens 349
503
tennesseense 349
var. nigrofastigiatum 349
tomentosum
var. Capitatum 350
var. compactum 349
verruciferum 349
var. surreptans 349
verruculigerum
var. formosanum 350
var. verruculigerum 350
vesuvianum
var. nodulosum
f. umbricola 345
f. verrucosum 350
vimineum 350
weberi 350
Sticta
lacera 399
oregana 350
Stioclettia 422
luzulina 422
Strangospora
senecionis 350
Strigula
submuriformis 299
Sulcaria
badia 350
isidiifera 350
Talaromyces
flavus 364-365
Taphrophila 69
Taxomyces 74
andreanae 71-72, 73-75, 77-80
Tephromela
elixii 351
muscicola 351
nashii 351
septentrionalis 351
tropica 351
velloziae 351
Terfezia
arenaria 209
berberiodora 210, 215
cadevalli 210
claveryi 210
hispanica 210
leonis 209
leptoderma 210
olbiensis 210
oligosperma 212, 215
pallidum 210
Thelidium
504
[Thelidium] umbilicatum 315
Thelomma
mammosum 296
Thelotrema
brasiliana 351
Thraustotheca 183-190, 192
achlyoides 186-188
caucasica 187, 191-192
clavata 183-190, 192
irregularis 188
primoachlya 183, 186-189
unisperma 186, 188
var. litoralis 188
var. unisperma 188
Thyronectria 43
balsamea 43
berolinensis 43
lamyi 43
Tiarosporella
paludosa 150
Toninia
arctica 351
bullata 352
ruginosa
subsp. pacifica 352
var. andicola 352
tristis
subsp. canadensis 352
subsp. pseudotabacina 352
Toxosporiopsis 65
Trametes
malicola 4
Trapeliopsis
hainanensis 352
pseudogranulosa 352
Trappea 215
darkeri 210
Trechispora
farinacea 407
Trematosphaeria 415-417
pachycarpa 415-418
Trichaptum
abietinus 16
biforme 16, 19-20
Tricholomopsis 385
Triparticalcar 292
arcticum 289
Tuber 93-97, 201
aestivum 93-97, 99-100, 210
asa 211
borchii 210, 215
brumale 211
excavatum 211
ferrugineum 212
foetidum 211
gennadii 201, 211
levissimum 201, 211
lutescens 210
maculatum 201, 211
malenconii 201, 211
maresa 211
melanosporum 94, 96-97, 211, 213
mesentericum 95, 97, 211-212
multimaculatum 201, 212, 214
nitidum 212
oligospermum 212
panniferum 212
puberulum 212
rufum 212
sinuosum 204
uncinatum 93-94, 96
Tubercularia
chaetospora 62-63
Tubeufia 67, 69
trichella 67-68
Tuckermannopsis 395-396
americana 396, 398
aurescens 396
aokesiana 396
chlorophylla 397, 401-402
ciliaris 397
coralligera 397
fendleri 398
gilva 398
hepatizon 398
merrillii 396, 398-402
microphyllica 399
oakesiana 400
orbata 396-397, 400
pallidula 400
platyphylla 400
platyphylloides 400
sepincola 401
ulophylloides 402
Tylopilus 374, 377
Tyromyces
fragilis 10
fumidiceps 16-17
galactinus 16-17
humeana 16-17
semipileatus 9
Umbilicaria
badia 314
lambii 352
phaea 352
Urceolella 178
505
[Urceolella] aasii 177-178 tubulosus 396, 402
crispula 178 viridis 402
tuberculiformis 178
Urohendersonia
pongamia 150 Wawea
Usnea fruticulosa 355
acromelana 352
durietzii 353
eulychniae 353 Xanthoparmelia 129
kuehnemanrii 353 austroconstrictans 101, 120-122
neoguineensis 353 canobolasensis 101, 122-123, 125
neuropogonoides 353 chlorochroa 328
nidularis 353 constrictans 120, 122
perpusilla 326 cylindriloba 355
pseudoceratina 353 glareosa 122-123
tanzanica 353 hyposalazinica 101, 123, 125
trachycarpa 326 kiboensis 316
Ustulina 233 lipochlorochroa 355
masonii 101, 124-125
mollis 356
Verrucaria mougeotina 124
bagliettoi 353 multipartita 124, 126
bisagnoensis 353 reptans 330
boccana 354 rubrireagens 122
contardinis 354 subdistorta 331
diplotommoides 354 subnuda 123-124
durietzii 354 sulcifera 331
f. rhabdota 354 trirosea 101, 124-126
fuscella victoriana 124
f. omblensis 354 xanthofarinosa 101, 127
gorzegnoensis 354 xanthosorediata 101, 129
haeyrenii 354 Xanthopeltis
"ha{umlat]yrenii" 354 rupicola 356
imperfecta 354 Xanthopsorella
incompta 354 texana 337
lacustris 354 Xanthoria
langhensis 355 candelaria 323, 335
latebrosoides 355 filsonii 356
omblensis 354 fulva 324
ornata 355 polycarpa
pseudomacrostoma 355 var. maritima 356
putnae 355 sorediata 304
rapallensis 355 Xylaria 228, 254
santensis 355 Xylocladium 219, 229, 233, 241, 243,
savonensis 355 247
Verticladiella 411-412
Vezdaea
rheocarpa 355 Zahlbrucknerella
Vuilleminia 85 patagonica 356
Vulpicida 395, 403 Zelleromyces
canadensis 397 stephensii 157
juniperinus 399
pinastri 397, 400
tilesii 401-402
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507
ERRATA, VOLUME FORTY-SIX
Pages 437-440 for.) Lema read Timah
ERRATA, VOLUME FORTY-SEVEN
Page 102 for _ P. subtabacina read B. subtabacina
PUBLICATION DATES
MYCOTAXON Volume 46: 1-508 (January-March)
was published on February 4, 1993
Correction:
MYCOTAXON Volume 45: 1-548 (October-December)
was published on October 30, 1992
508
Reviewers, Volume Forty-Seven
The Editors express their appreciation to the following individuals who have, prior to acceptance for
publication, reviewed one or more of the papers appearing in this volume:
T. Ahti
J. Ammirati
A. W. Archer
allan
ec) ‘Chang
Ef Grane
. Fogel
. Guarro
. Hawksworth
R. A. Humber
I. Karnefelt
J. W. Kimbrough
R
R,E
D. M. Langsam
G.-z Ma
D. Malloch
G. Morgan-Jones
G. M. Mueller
A. Nawawi
D. E. Padgett
O. Petrini
E. Punithalingam
A. Rambelli
L. Ryvarden
E. Sérusiaux
A. Strid
F. A. Uecker
G. Van der Westhuizen
S. G. Vanev
D. Verdon
M. A. Vincent
C. Walker
R. Watling
C. B. Wolfe
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